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DEPARTMENT for ENVIRONMENT, FOOD and RURAL AFFAIRS CSG 15 Research and Development Final Project Report (Not to be used for LINK projects)

Two hard copies of this form should be returned to: Research Policy and International Division, Final Reports Unit DEFRA, Area 301 Cromwell House, Dean Stanley Street, London, SW1P 3JH. An electronic version should be e-mailed to [email protected]

Project title Agricultural practice and bats: A review of current research literature and management recommendations

DEFRA project code BD2005

Contractor organisation Bat Conservation Trust and location 15 Cloisters House 8 Battersea Park Road London SW8 4BG

Total DEFRA project costs £ 12,225.00

Project start date 01/02/03 Project end date 03/09/31

Executive summary (maximum 2 sides A4)

 Agricultural intensification processes are widely-reported to have been detrimental to farmland wildlife. Although the impacts of agricultural management practices on birds and other groups have been widely studied, their effects on bats are less understood.

 A review collated scientific literature on the occurrence of bats in agricultural landscapes and the impact of management on bats, their key habitats and invertebrate prey. Data on four aerial-insect feeding birds were also collated.

 All British bat species occur in agricultural landscapes to some extent. Information on occurrence and habitat utilisation (in the context of agricultural landscapes) was obtained for eleven species.

 Studies on bats were primarily conducted in lowland landscapes dominated by grassland or mixed farming. There was a paucity of data on the occurrence and behaviour of bats in the uplands and in lowland arable landscapes.

 The review drew on a smaller pool of studies than would have been available for most farmland birds. The majority of studies focused on describing basic species ecology.

 The relatively small pool of studies restricts the present understanding of intraspecific variation in factors such as diet, foraging behaviour, home range etc.

 In the main, research addressed the distribution and activity of foraging and commuting bats, with few publications focusing on roosting or hibernation behaviour.

CSG 15 (Rev. 6/02) 1 Project Agricultural practice and bats: A review of current DEFRA BD2005 title research literature and management recommendations project code

 Much bat activity was associated with the non-crop component of agricultural landscapes (i.e. woodland, water and riparian habitats, hedgerows and linear elements). Woodland and water were important foraging habitats. Hedgerows and linear features were used by several species for commuting between foraging and roosting areas.

 Among production habitats, arable fields were widely reported to be avoided. However a recent study has reported natterer’s bat foraging over arable fields and common pipistrelle is known to feed across many habitats, including arable.

 A number of species foraged over agricultural grasslands, including several for which invertebrates associated with cattle dung form an important constituent of diet. The relative value of unimproved, semi-improved and intensively managed grassland was not conclusively resolved with findings conflicting among species and studies.

 Important items in the diet were Diptera, Lepidoptera and Coleoptera. Information on the habitat requirements of families listed in diet reviews, and on the predicted impact of agricultural management practices was collated.

 It is widely believed that agricultural intensification practices have contributed to the decline of several European bat species. Proposed mechanisms include loss of roost sites, degradation or reduced accessibility of foraging areas, depletion of prey and direct / indirect exposure to pesticides.

 The main research effort has focused on the potential impact of reduced food abundance on bat activity. Evidence that aerial insects may be a limiting resource is supported by studies of swallows in Denmark, where reduced clutch sizes, poorer quality nestlings and reduced abundance of adults were associated with low densities of aerial insects.

 Long-term studies provide evidence that several of the invertebrate groups important in bat diet including moths, craneflies and dung beetles have declined in agricultural landscapes.

 Recent studies investigating the activity of bats in different farming systems have found higher levels of activity on organic than conventional land and have attributed this to higher overall insect abundance of key diet insects on organic farms. More general studies on invertebrates have been equivocal, with variations among groups. Some differences are attributed to patterns of landscape heterogenity rather than management differences per se.

 Responses to individual farm management practices varied among taxonomic groups but pesticides including insecticides and herbicides were reported to have been detrimental to many bat-food insect groups through direct toxicity or depletion of plant foods or microhabitat. Endo-parasite control treatments are known to affect dung fauna, particularly dung flies and dung beetles, but the effects on higher trophic levels has not been quantified at landscape level.

 Mechanical practices including soil cultivation, drainage, cutting and grazing affect the abundance and species composition of assemblages of bat-food insects. Diptera are particularly sensitive to drainage and changes in vegetation structure along ditches, while the soil-dwelling larval stages of Diptera and Coleoptera may be destroyed as part of managements that disturb the soil. As many of these operations occur as a normal part of countryside management, management for spatial and temporal heterogeneity is most likely to support biodiversity.

 Existing agri-environment schemes contain a variety of prescriptions for habitats of key importance to bats including water and riparian features, trees and woodland, hedgerows and grassland. Few prescriptions specifically target mammals.

 Agri-environment advisors were in most cases aware of generalised habitat requirements of bats, but placed more emphasis on the importance of hedgerows and linear habitats than any other landuse type. Bats were often used to illustrate the importance of habitat management, rather than targeting management specifically for bats.

 Lack of local fine-scale data on species distributions was the most commonly cited obstacle to implementing conservation measures for bats on farmland.

CSG 15 (Rev. 6/02) 2 Project Agricultural practice and bats: A review of current DEFRA BD2005 title research literature and management recommendations project code

Scientific report (maximum 20 sides A4)

CSG 15 (Rev. 6/02) 3 Project DEFRA title project code

Agricultural practice and bats: A review of current research literature and management recommendations

1. Introduction

1.1 Rationale for considering the impact of agricultural practice on bats

Agricultural land can be a diverse habitat for wildlife, and the substantial area that this land-use represents within the U.K. and within Europe has the potential to create areas rich in farmland species (Sotherton, 1998). During the latter half of the twentieth century, however, many regions underwent agricultural intensification processes that are now widely-known to have had serious consequences for the general environment (Skinner et al., 1997) and for biodiversity e.g. (Stoate et al., 2001; Benton et al., 2002; Robinson & Sutherland, 2002; Brereton, In press). Reviews such as (Robinson & Sutherland, 2002) outline the complexity of the processes that have occurred and the breadth of impacts, some of which are probably still insufficiently recorded. Intensification processes have occurred across all sectors of agriculture (i.e. arable land, grassland, horticultural crops). Examples of intensification are the increased use of inorganic fertiliser, wide-scale application of pesticides, removal of boundary features such as hedgerows resulting in large fields and coarse-grained landscapes, land drainage, and the exchange of small-scale mixed farming systems for larger, specialist farms. Considerable effort is now being directed towards mitigating the impact of agriculture on the environment, the development of sustainable farming, and the restoration of habitats and species that were previously depleted.

Although precise population statistics are difficult to achieve (Altringham, 2003), particularly in a historical context, it is widely considered that many bat species underwent serious population declines during the twentieth century, both in Britain and across western-central Europe (Schober & Grimmberger, 1989; Altringham, 2003). Evidence of these changes includes range contractions of well-documented species and reduction in colony size / colony disappearance from traditional roosting sites. Factors contributing to these declines extend beyond agriculture and include the use of toxic wood treatment chemicals in roof spaces, disturbance and persecution, and unintentional destruction of roosts. Agriculture has, however, by virtue of controlling the management of more than 76% UK land surface (Robinson & Sutherland, 2002), including many features closely associated with bats, the potential to impact severely on UK populations. In the light of the negative impacts of past agricultural intensification on many farmland birds, wildflowers and invertebrates, it is reasonable to assume that a group of mammals with complex life-histories and reliance on an insect diet would also be vulnerable to intensive farming practices.

All bats in the UK are protected, and recent legal obligations to reverse declines and maintain populations of species assigned Biodiversity Action Plan (BAP) status have drawn attention to the need to identify and manage positively those factors that impact populations. Due to the difficulty of working on small nocturnal highly mobile flying animals, ecological field studies on bats are, in comparison to birds, a recent development. Continuing technical advances (in bat detectors, radio-tracking transmitters, night vision equipment etc) are opening this field at a time of increasing scientific and public interest. The extent to which existing ecological literature on bats and their requirements could be used to inform policy for agricultural land was unknown at the time of this review, providing an opportunity to influence future research and conservation strategy. The latter aim is timely, since the review coincides with a revision of the major agri-environment schemes in England.

The case for species as indicators of environmental health (bio-indicators) has been made for many organism groups. The term seems particularly apt for bats, which have complex habitat requirements, including different roosts for hibernation, mating and rearing young, exhibit a high degree of site fidelity, are reliant on a variable invertebrate food-resource and are sensitive to factors at scales from micro-climate to landscape. As technical advances increase the accuracy with which species distribution and abundance can be assessed, the case for using bats as measures of sustainable management is likely to strengthen.

1.2 Scope of review

This review collates ecological research examining bat activity in agricultural landscapes or in wider landscapes that incorporate an agricultural component. The review is restricted to resident British bat species, but includes data collected in continental Europe where this examines habitats and farming systems similar to those encountered in Britain. In a parallel discipline, ornithology, research has addressed a suite of species collectively known as “farmland birds”, typically granivorous passerines or game-birds of arable habitats, but also waders associated with mixed / grassland farms. The term “farmland bats” is less defined at this time as the basic ecology of the majority of species is still being determined and, in contrast to birds, a larger proportion of the British species pool is considered

4 Project DEFRA title project code to occur on farmland to some extent. The policy of this review has therefore been to collate data on any bat species referred to in an agricultural context. BAP species (see Appendix 1) were considered a priority, but again the review was not restricted to these since it was anticipated that the pool of available data might be insufficient. Instead, all species were considered, in order to examine common/contrasting responses to land use. Greater horseshoe bats, the most widely studied of British bats, are not a prime focus of this review, because they are already the subject of detailed conservation plans (Ransome, 1996; Ransome, 2000) and implementation programmes (Longley, 2003). The review’s main role was to collate the responses of less studied species to agriculture and has the advantage of expanding the geographical context beyond southern and western Britain where greater horseshoes occur. Trends in agricultural management (i.e. dominant crops, degree of intensification) vary along east-west transects, and along altitude gradients and are likely to exert different pressures on bats accordingly.

The review aims to collate evidence of direct agricultural impacts on bats or indirect impacts mediated, for example, through changes in the supply of invertebrate prey. The habitats and invertebrate groups evaluated are those identified as important to bats in the scientific literature. British and European materials on habitats and invertebrates were prioritised, although occasional reference is made to alternative sources where there is a paucity of more local information.

In recognition of a potential lack of historical information on bats in farmland habitats, and to examine parallels with other groups, the review was expanded to consider several birds with ecological requirements in common with bats. The species selected were swallow Hirundo rustica, house martin Delichon urbica, swift Apus apus and spotted flycatcher Muscicapa striata. Swallows, house martins and swifts feed on aerial invertebrates by hawking and diets overlap with several bat species (section 3.2). They are central place foragers, which breed colonially on buildings. Swifts and house martins nest in more urban surroundings, while swallows are more closely associated with rural villages and farmed landscapes. Spotted flycatcher is a bird of woodland edges, which makes short flights from perches to catch aerial prey or in poor weather gleans or flushes insects from trees. Unlike British bats, these species are long-distance migrants, with survival also influenced by conditions at their African wintering grounds, and along migration routes. Also in contrast to bats they are relatively short lived with the potential to rear several young in a season. Though potentially affected by similar factors to bats in their summer ranges, they might therefore be expected to exhibit different rates of population change.

For the purpose of this review, agricultural habitats are considered to include arable, grass and horticultural crops, non-crop habitats (e.g. hedgerows, shelter-belts, ponds) and farm buildings. Detailed aspects of woodland management, where forestry or a non-agricultural use are the prime aim are considered beyond the scope of this review, although some reference is made to the creation of farm woodlands, since these are promoted by schemes that target farmers.

The review is a scoping exercise, to identify key patterns in agriculture with potential to impact bat populations, identify research gaps and promote good practice by making research findings more available to those that implement conservation on agricultural land. To facilitate this latter aim, the review collates basic information on management options for habitats important to bats in agri-environment schemes, and investigates the degree to which bat conservation is already promoted on farms, by discussing conservation approaches with prominent networks of agri-environment advisors. The main purpose of the review is therefore to bring together information on research findings and current conservation practices and by doing so, improve bat conservationists’ understanding of agricultural issues, and bring new bat research to the attention of policymakers and agri- environment advisors.

2. Methodology

2.1 Literature review

2.1.1 Electronic searches

Extensive literature searches were carried out using several major web based scientific databases. The majority of searches were done in ISI Web of Science: Science Citation Index (Expanded), which contains over 17 million references from 1981 to present, and CABdirect, which contains over 4 million records running from 1973 onwards. Further searches were carried out in a number of e-journal databases, mainly: Science Direct, JSTOR and SwetsWise.

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A wide range of keywords and wildcards relating to bats, agricultural land and invertebrates important to bats were used during the searches. Some examples of the main keywords are given below. A variety of combinations of these were used during the searching process. Additional sources were then compiled from the reference lists of key papers.

Table 1. Search terms used in electronic searches for reference material.

Bat related Agriculture related Invertebrate related Bats Farm* Diptera or fly Chiroptera Agri* Nematocer* Microchiroptera Pastur* Chironomid* Diet Grass* Tipulid* Foraging Horticult* Lepidoptera or moth or butterfly Pipistrell* Pesticide* Coleoptera or beetle Insecticide* Aerial or flying * Wildcards used where words can have several endings

2.2 Collation of agri-environment scheme materials

Up to date agri-environment scheme literature was obtained from relevant government departments for England, Wales, Scotland and Northern Ireland. Although Wales, Scotland and Northern Ireland fell outside the administration of DEFRA, the funder of this review, these schemes were considered to be of complementary interest, since they operate within similar habitat and climate zones. Features peculiar to these schemes could be considered for adoption in England where they appear to bring additional benefits for bat conservation.

Due to time constraints, the review considered those habitat prescriptions likely to impact most on bats for cross- comparison among schemes. Four broad habitat categories deemed important to bats were selected after reference to scientific literature. These were: hedgerows, grasslands, waterbodies (including their margins) and trees. Summaries of management prescriptions relating to these categories were produced for all schemes. In addition, prescriptions detailing regulation of pesticide usage were collated, in recognition of potential indirect impacts on bats through their invertebrate prey.

2.3 Discussions with agri-environment advisors

A series of discussions were held with persons engaged in different aspects of agri-environment advice ranging from first point of contact with farmers and land managers through to development and administration of agri- environment agreements. These discussions aimed to: improve bat conservationists’ understanding of the process by which conservation measures are encouraged on farmland; determine the extent to which advisers currently target bats in their recommendations; recognise potential opportunities to assist in the provision of better advice; identify particular obstacles to the implementation of practices that would support bats; gain examples of local initiatives and good practice and identify what future resources would be most appreciated by agri-environment personnel.

To obtain information relating to England discussions were held with 12 FWAG advisors, 7 RDS regional biodiversity co-ordinators / ecologists, and 1 RDS project officer. Contact lists were supplied by the RDS technical advice unit and FWAG central office and were selected at those offices to give broad geographical coverage across England, the area addressed by DEFRA agri-environment schemes (Countryside Stewardship and ESA). In addition, 2 FWAG advisors and 3 SAC conservation advisors working in Scotland and 1 FWAG advisor and 1 CCW Tir Gofal officer working in Wales were contacted in order to identify alternative approaches facilitated by other schemes that might be suitable for adoption in the area administered by DEFRA. Since the objective was to gain opinions from a representative cross-section of advisers, no attempt was made to target those with specialist knowledge of bats, although a number of the advisers we spoke to held English Nature bat conservation licenses. Following recommendations from this initial sample of advisers, contact was made with a small number of individuals with extensive experience of conserving bats on agricultural land.

Structured discussions were held lasting approximately 10-20 minutes during which geographical coverage, the main types of farmland encountered in the adviser’s region and the process of recognising wildlife habitat on farmland and planning for biodiversity were explored. Interviewees were asked whether bats were often mentioned

6 Project DEFRA title project code during discussions with land managers (by either the adviser or land manager), and were asked to describe the types of recommendations given if advice was made with bats in mind. Advisers were asked to identify prescriptions in agri-environment schemes that could be used to support bats and whether they encountered particular obstacles to putting conservation measures that would support bats in place. General information on the most popular prescriptions in the advisers’ area was collated and where there was evidence of work targeted at bats, examples for use as potential case studies were requested.

While best efforts were made to ensure a broad range of individuals were contacted it should be emphasised that this exercise was a discussionary process, the intention being to gain general insights into the conservation focus on agricultural land and not to conduct a statistical analysis.

3. Results

3.1 General occurrence of bats on farmland and association with habitat features

During the review several types of study were encountered from which information relating to bats and agriculture could be derived. The majority of studies focused on the relative distribution / activity of bats at landscape scale based on detector studies or radio-tracking tagged individuals. These studies recorded the occurrence of bats in different landscapes in relation to land-use type and component features, to determine the basic ecology of individual species. Key results of such studies are summarised in Appendix 2 (bat detector studies) and Appendix 3 (radio-tracking studies) where species are studied in predominantly agricultural landscapes, or wider landscapes that included comparison of bat occurrence in agricultural habitats with other land-uses. The most sophisticated studies took into account the relative amounts of different land-types / habitat features in the study area, allowing the significance of any association to be determined. A small number of the studies summarised in Appendices 2 and 3 used these techniques to make comparisons between a very limited number of feature types e.g. new woodlands with arable (Moore et al., (submitted)), smooth or rippled streams (Warren et al., 2000).

In general the studies using bat detectors evaluated larger study areas, sometimes at national scale e.g. (Walsh & Harris, 1996a; Walsh & Harris, 1996b), although they were occasionally used to examine the movements of bats from a single colony. Owing to the difficulty of distinguishing between species that make similar sounds, detector studies often refer to species groups e.g. Myotis-Plecotus group. Radio-tracking studies usually follow individuals from a colony to foraging areas, enabling home-range, distance travelled in a night and use of commuting/foraging habitat or night-roosts to be recorded. Such studies have the advantage of being able to separate sex or age differences in behaviour, but until very recently, tags of sufficiently small size were not available and this has resulted in a very small pool of studies for comparison. For this reason, behaviours described for individual species are typically based on small sample sizes.

A few papers compared specific aspects of farm management based on pairs of matched habitats in different farming systems, or as a manipulative experiment. These studies, and those that evaluated hypotheses to explain the preference for certain features are considered separately.

The studies listed in Appendices 2 and 3 do not represent UK regions equally, reflecting in part the locations of specialist bat research groups. Three papers (Walsh & Harris, 1996a; Walsh & Harris, 1996b; Russ & Montgomery, 2002) are unusual in surveying at a national scale. In general, studies had a pronounced southern and westerly focus (i.e. south-west England, Wales, southern England, Ireland). Scottish habitat studies focused on north-east Scotland. Southeast England, a region where agricultural intensification has been extreme was represented only by a study of a serotine colony in Cambridgeshire (Robinson & Stebbings, 1997) and a Leisler’s colony in Kent (Waters et al., 1999). In terms of agricultural landscapes, predominantly arable landscapes were represented only as part of a national distribution survey (Walsh & Harris, 1996a; Walsh & Harris, 1996b) and by the Cambridgeshire serotine study (Robinson & Stebbings, 1997). Most of the studies in Appendices 2-3 were conducted in lowland pastoral e.g. (McAney & Fairley, 1988; Vaughan et al., 1997; Shiel & Fairley, 1998; Shiel et al., 1999; Motte & Libois, 2002; Schofield et al., 2002) or lowland mixed farming landscapes e.g. (Racey & Swift, 1985; Catto et al., 1996; Entwistle et al., 1996). Upland landscapes were represented by a single behaviour study (Warren et al., 2000) and this addressed distribution along a watercourse rather than on agricultural land per se. Volunteer safety and low density of surveyors restricted the coverage of upland habitats in a national bat distribution survey (Walsh & Harris, 1996a; Walsh & Harris, 1996b).

Despite the gaps and potential biases in UK coverage, information on the occurrence and habitat selection in agricultural landscapes was collated for eleven of the 16 resident UK bat species (Appendices 2-3). No data on

7 Project DEFRA title project code occurrence or habitat selection within the context of agriculture were collated for Bechsteins, barbastelle, Nathusius pipistrelle, grey long eared or Brandt’s bat. This is more likely to be due to the lack of studies on these species, and the fact that several of these (e.g. grey long-eared, Bechsteins) are very rare than to indicate these species complete avoidance of agricultural habitats. (Wickramasinghe et al., In press) noted that all UK bats occur within agricultural landscapes, and recorded 14 species during a study of organic and conventional farms located across seven counties in England and Wales. (Moore et al., (submitted) reported 7 species within the smaller study area of the lower Derwent valley in East and North Yorkshire. (JNCC, 2001) lists features closely associated with agricultural management as priority habitats for all six UK bats with BAP status (Appendix 1), including barbastelle and Bechsteins.

At a coarse-scale, agricultural land may be divided into habitats associated directly with production (e.g. arable land, grassland) and non-crop (e.g. woodland, hedgerow, watercourse). Among the production habitats, grassland was often categorised at a finer resolution (e.g. unimproved, improved, cattle-pasture etc) but arable land was not resolved into different crop types. Non-cropped habitats were frequently described more finely (e.g. broad-leaved wood versus coniferous wood). A clear majority of studies reported that bats avoided arable land. This finding was reported for overall bat activity (Walsh & Harris, 1996a; Walsh & Harris, 1996b), brown long-eared (Moore et al., (submitted)), leislers (Waters et al., 1999), (Robinson & Stebbings, 1997), lesser horseshoe (Bontadina et al., 2002; Motte & Libois, 2002) and greater horseshoe (Duvergé & Jones, 2003). Although this trend would appear to be almost universal, there is recent radio-tracking evidence of natterer’s bat, a species usually associated with woodlands, preferentially foraging over arable crops (Aegerter, pers. comm). Another species, common pipistrelle has been reported to feed evenly across all lowland habitats including arable (Vaughan, 1997) and to forage around mature trees overhanging arable land (Davidson-Watts, unpublished).

The value of agricultural grassland to bats is less clear as findings vary among species and among studies. Across the UK, (Walsh & Harris, 1996a; Walsh & Harris, 1996b) reported that lowland unimproved grassland was the only grassland habitat not consistently avoided and that semi-improved and improved grassland sites were among those habitats least used by bats. Other authors e.g. (Vaughan et al., 1997) considered the techniques used in the Walsh survey to have been most sensitive to pipistrelle species, but the results are supported in part by other research where resolution to species-level has been possible. For example, another detector survey (Vaughan et al., 1997) recorded high levels of Myotis-Plecotus activity over unimproved grassland and improved cattle pasture, and common pipistrelle activity in unimproved and amenity grassland and cattle pasture (as well as a diverse range of non-crop habitats). The relative value of unimproved, semi-improved and improved grassland to bats has not been clearly resolved, however, as there are also reports of bats utilizing improved grassland in preference or equivalence to more extensive management. For example, noctule and leisler activity was high over improved pasture in southwest England (Vaughan et al., 1997) but in Ireland Leisler’s appeared to avoid improved grassland (Russ & Montgomery, 2002).

Several species were reported to forage selectively over pasture e.g serotine (Catto et al., 1996; Robinson & Stebbings, 1997), Leisler’s (Shiel et al., 1999; Waters et al., 1999), greater horseshoe (Duvergé & Jones, 2003), and noctule (Vaughan et al., 1997; Wickramasinghe et al., In press). None of the studies examined set out to discriminate between pastures grazed by different animals as a main purpose of the study, but use of cattle-grazed grassland was often reported. In southwest England a colony of greater horseshoe bats preferred cattle-grazed pasture to sheep or horse-grazed land (Duvergé & Jones, 2003). Published studies on lesser horseshoes have previously emphasised their avoidance of open habitats such as arable and grass fields and the use of sheltered linear features as commuting routes to woodland feeding areas (McAney & Fairley, 1988; Bontadina et al., 2002; Motte & Libois, 2002; Schofield et al., 2002). However, recent observations by Wray (pers. comm.) suggest that the presence of cattle may increase the attractiveness of pasture for this species. Although largely untested in the bat literature, other insectivorous species with similar ecological requirements to bats are known to make similar fine- scale discrimination between habitats. For example, the presence of livestock on grassland farms was the most efficient predictor of the presence of breeding swallows (Ambrosini et al., 2002). Swallow researchers have attributed this effect to the impact of cattle on prey availability. (Moller, 2001) found the abundance of flying insects to be significantly higher in the presence of livestock. Two mechanisms may be involved since cattle dung is food for a complex invertebrate fauna, and animal movement may result in insects becoming more available to aerial predators when insects are flushed. Hirundines are able to exploit disturbed prey, taking much food from around the feet and noses of grazing cattle (O'Connor & Shrubb, 1986). Bats are known to respond rapidly to short- term changes in habitat quality. For example, both serotines (Catto et al., 1996) and noctules (Haysom, personal observation) have been observed hawking for insects over recently mown grass.

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All the studies examined identified non-crop habitats to be important foraging habitats for bats. Across the UK, (Walsh & Harris, 1996a) reported selection of woodland and water habitats and edge / linear features such as tree- lines, hedgerows and rivers by bats. Overall bat activity and that of various species was particularly associated with the presence of water. For example, in south-west England, 70% of all bat passes recorded by (Vaughan et al., 1997) occurred near rivers or lakes. Species for which water was a preferred foraging habitat included daubenton’s bat (Swift & Racey, 1983), a combined Myotis/Myotis-Plecotus group (Vaughan et al., 1997; Russ & Montgomery, 2002), common pipistrelle (Russ & Montgomery, 2002), soprano pipistrelle (Vaughan et al., 1997; Russ & Montgomery, 2002) Davidson-Watts, unpublished data); whiskered (Moore et al., (submitted)), noctule (Vaughan et al., 1997; Wickramasinghe et al., In press) and Leisler’s bat (Shiel et al., 1999; Russ & Montgomery, 2002). One of the four lesser horseshoe studies reported intense foraging activity over water-bodies (McAney & Fairley, 1988), and a second study the use of riparian woodland (Schofield et al., 2002). Physical and chemical characteristics of water-bodies appear to influence their use. Foraging daubenton’s and common pipistrelle bats preferred stretches of smooth water with trees on both banks (Warren et al., 2000). Rapid water was thought to impede echolocation of prey, and densities of aerial insects were higher on river stretches with trees. Overall bat activity was reduced by 11% and prey capture attempts by 28% in polluted water (Vaughan et al., 1996). The activity of both pipistrelle species was reduced in a polluted area, while serotine, Nyctalus and Myotis activity was not significantly affected. There was some evidence, however that the foraging efficiency of Myotis improved in polluted water.

Woodland or trees were listed as key foraging habitats for all the species encountered. The degree to which discrimination was made between continuous blocks of habitat, through to tree-lines and individual trees, varied among the species. There were contradictory reports on the comparative importance of woodland core and edge habitats. (Walsh & Harris, 1996a; Walsh & Harris, 1996b) recorded greater activity at woodland edges than centres while (Russ & Montgomery, 2002) found use of edge versus centre habitats varied among species and (Vaughan et al., 1997) reported no significant difference in overall bat activity between the two locations. In studies of new farm woodlands (Moore, 2002; Moore et al., (submitted)) found that blocks of establishing saplings were more attractive foraging areas for all bat species combined and pipistrelle species than adjacent arable land. Where tree species was considered, there was a general preference for broad-leaved over mixed or conifer woodland (Walsh & Harris, 1996a; Russ & Montgomery, 2002) although bats including common pipistrelle, soprano pipistrelle, Leisler’s and Myotis were recorded in mixed or coniferous plantation (Vaughan et al., 1997; Russ & Montgomery, 2002). Deciduous, mixed and coniferous woodland were all listed as preferred foraging habitats for brown long-eared bats (Swift & Racey, 1983; Entwistle et al., 1996) and studies of lesser horseshoe bats in Belgium and Germany place them in deciduous, mixed and conifer forests (Holzhaider et al., 2002; Motte & Libois, 2002). Greater horseshoe bat activity was reported higher in deciduous than coniferous woodland (Duvergé & Jones, 2003).

Hedgerows or trees associated with hedgerows were named among the preferred foraging habitats of whiskered bats (Moore et al., (submitted)) and brown long-eared bats (Moore et al., (submitted)), but (Russ & Montgomery, 2002) reported that hedgerows were among those features least used by various species in a Northern Irish survey. High levels of bat activity were reported associated with hedgerow habitats in the national UK survey (Walsh & Harris 1996b) and, in other studies, were often reported as features that facilitated commuting, providing foraging opportunities on route to primary foraging areas e.g. (Motte & Libois, 2002). Studies in the Netherlands (Limpens & Kapteyn, 1991; Verboom & Huitema, 1997; Verboom & Spoelstra, 1999) considered the role of linear landscape elements including hedgerows in connecting other habitats and the mechanisms behind their selection. Hypotheses evaluated have included provision of shelter from wind leading to reduced energy consumption; provision of richer supplies of insects, avoidance of predation and range of echolocation. In relation to shelter, (Limpens & Kapteyn, 1991) reported that linear elements less than 1m high were seldom used in contrast to elements with tall vegetation, and that bats tended to fly along the leeward side of linear features, where highest densities of aerial insects also occurred. (Verboom & Spoelstra, 1999) found that common pipistrelles in an agricultural landscape focused their activity closer to tree lines at high wind speeds or angles of wind incidence from 45 o to 95o. Small species such as pipistrelle sp. were thought to be more dependent generally on the presence of linear features than large species e.g. serotine which frequently crossed open ground (Verboom & Huitema, 1997). The use of linear features as commuting routes is not confined to small bats however. Greater horseshoe bats preferred flying within 5m of field boundaries (on average flying within 2m) and flew along the side of hedgerows rather than above them (Duvergé & Jones, 2003). Serotines were also recorded travelling along tall hedges and tree-lines (Robinson & Stebbings, 1997). Comparable ornithological research has addressed the use of hedgerows by swallows in agricultural landscapes (Evans et al., 2003). Swallows selected vegetated boundaries in bad weather and there was a non-significant trend towards selection in good weather, the habitats selected being those with highest food availability.

Data summarised in Appendices 2 and 3 show that all species were reported to feed in more than one habitat type, and that the number of habitat types utilised varied among the species. Common pipistrelle was reported to be

9 Project DEFRA title project code particularly broad in its utilisation of habitat for foraging (Davidson-Watts, unpublished; (Vaughan et al., 1997; Russ & Montgomery, 2002). The relative use of different habitat types was reported to vary seasonally in serotine (Robinson & Stebbings, 1997) and greater horseshoe (Duvergé & Jones, 2003). Radio-tracking studies (Appendix 3) provide further information on habitat use at individual and colony level. Individual bats (of different species) varied in the number of feeding locations used in a night and in the patterns of feeding location utilisation (i.e. visited once/many times in a night). Brown long eared bats (Entwistle et al., 1996; Moore et al., (submitted)) and pipistrelles ((Racey & Swift, 1985) Davidson-Watts, unpublished ) were loyal to individual feeding sites, returning to the same locations night after night but greater horseshoes shift the locations of foraging areas, resulting in changes in the area used over time (studies cited in (Duvergé & Jones, 2003)). Foraging pattern also differed between adults and juveniles in some studies. In contrast with adult brown long-eared, juveniles foraged in different locations each night (Moore et al., (submitted)).

Maximum direct distances travelled by individual bats between roost site and a foraging area ranged from 1.1 km for brown long eared (Swift & Racey, 1983) and 1.2km for lesser horseshoe (Motte & Libois, 2002) to 13.4km for Leisler’s (Shiel et al., 1999). Where several studies of a species were available intra-specific variation in distance travelled to a foraging site could be large. For example, (Moore et al., (submitted)) found brown long-eared bats travelling up to 7.4km from the roost, over twice the distance reported in a scottish study (Entwistle et al., 1996). Distances travelled by breeding females between roost and foraging area were reported lower than pre-parturition, or in comparison to males e.g. (Racey & Swift, 1985; Entwistle et al., 1996; Bontadina et al., 2002).

Few of the studies encountered addressed bat requirements other than foraging and commuting habitat, in agricultural landscapes. There was, for example, little information on roosting sites. (Duvergé & Jones, 2003) reported that 31.8% of greater horseshoe roost sites recorded in their study were related to farm buildings and a further 43.2% were located on or near farmland. In a Scottish study area encompassing agricultural land, villages and small towns, (Jenkins et al., 1998) found all soprano pipistrelle roosts in buildings to be within 550m of a permanent water body. Furthermore, most roosts had some form of linear vegetation link to a woodland area, or were immediately adjacent to woodland. (Oakeley & Jones, 1998) reported comparable results for soprano pipistrelle maternity roosts located in an agricultural landscape in Wiltshire and Somerset. Significantly more water and continuous hedgerow with emergent trees was found within a 2km radius of roost sites than around random points. The length of water with hedgerow or woodland on at least one bank was also significantly greater around roosts. Similarly in the Avon valley (Davidson-Watts, unpublished) maternity roots of soprano pipistrelle were mainly located within the flood plain, allowing foraging access to water bodies lined with trees. For long-eared bats, buildings containing roosts were located closer to woodland and water, and were surrounded by a greater area of woodland within a radius of 0.5km than random houses (Entwistle et al., 1997). Brown long-eared roosts also had a number of structural and microclimate differences to random houses (e.g. older buildings, larger and more complex roof spaces, warmer temperatures). Traditional barns may be used as summer roosts by pipistrelle sp., brown long eared, natterer’s and occasionally by serotines (Briggs, 1995), particularly where barns are more than 100 years old, roofed with thick beams. In the case of barns nothing is published regarding the additional habitat factors that may enhance the value of the buildings for these species.

Several of the radio-tracking studies encountered reported the use of night-roosts (e.g. in trees or buildings) by bats. Use of night-roosts was described for lesser horseshoes (Bontadina et al., 2002; Schofield et al., 2002), Leisler’s (Shiel et al., 1999), serotine (Catto et al., 1996), and greater horseshoe bats (Ransome, 2000; Duvergé & Jones, 2003). Night-roosts may be used for resting, grooming, eating large prey, or sheltering bats in poor weather (Shiel et al., 1999) and may be particularly important for pregnant bats, allowing them to range further from the maternity roost (Schofield et al., 2002).

Little information was encountered reporting the occurrence and use of hibernation sites on farmland. Bats such as natterer’s may be found hibernating in the hollow mortise joints of traditional farm buildings (Briggs, 1995) while (McAney, 1999) outlined the potential importance of caves and mines as hibernacula. Such sites have recently been discovered to be visited by large numbers of bats in the autumn for “swarming”, an activity that may be associated with mating (Altringham, 2003). Greater horseshoe bats remain the only species for which studies integrating all aspects of habitat use, including the use of hibernacula, have been attempted in the UK.

3.2 Utilization of invertebrate prey

All UK bats are insectivorous and the diet of fifteen species was the subject of a comprehensive recent review that collated data from 61 European studies (Vaughan, 1997). Data for all sixteen of the species confirmed resident in Britain are now available following the publication of (Barlow, 1997) which separated the diets of common and

10 Project DEFRA title project code soprano pipistrelle (grouped in other studies prior to recent developments in taxonomy). At the present time, data for the majority of species are drawn from a small pool of studies, differences in methodology restrict comparison and intraspecific variation in diet requires further investigation (Vaughan, 1997). In addition to summarising the key prey groups utilized by each species, (Vaughan, 1997) drew attention to five foraging strategies previously distinguished for bats: fast hawking, slow hawking, trawling, gleaning and perch-hunting (fly-catching). Such strategies are related closely to the species morphology and echolocation characteristics, and determine the category of prey that may be taken. Species that deploy more than one strategy might be expected to vary the mode of foraging according to environmental cues, to optomise prey intake.

Vaughan’s review forms the basis of Appendix 4 which lists, in descending order of approximate importance, invertebrate families that occur in the diet of British bats. Dipteran and Lepidopteran families are dominant in the diet with Coleopteran families taken by many species, and a small number of other orders including Hemiptera (true bugs, plant hoppers and leaf hoppers) and Arachnida (spiders and their allies) taken in small quantities. Nine British bats eat mainly Diptera, and all species eat Lepidoptera to some extent (Vaughan, 1997). The reliance of British species on Diptera and Lepidoptera contrasts with the situation in North America and Africa, where Lepidoptera and Coleoptera are the predominant groups taken and it is postulated that the importance of Diptera may be related to Britain’s cool, damp climate (Vaughan, 1997). (Vaughan, 1997) identified several species with more specialist diets. Barbastelle and long-eared bats ate mainly moths and greater horseshoe took mainly beetles and Lepidoptera. For several species taxonomic resolution of prey identified through analysis of droppings or collected below feeding perches was sufficient to provide evidence of associations with particular habitats. Dietary evidence associated Bechstein’s bat with woodland Diptera and Lepidoptera (Vaughan, 1997), lesser horseshoe with damp woodland and water (Vaughan, 1997) and daubenton’s, Leisler’s and pipistrelle (particularly soprano pipistrelle) with aquatic insects (Barlow, 1997; Vaughan, 1997). Serotines specialise on beetles, particularly cockchafers and dung beetles, confirming the importance of cattle pasture as a feeding habitat for this species (Catto et al., 1994). Daubenton’s bat was reported to have the broadest dietary niche, taking insects in 16 taxonomic categories and lesser horseshoe, Nathusius pipistrelle and Leisler’s the most restricted diets (nine categories) (Vaughan, 1997). The degree to which bats select types or sizes of prey is unknown for the majority of species. Greater horseshoes are reported to take few flies and parasitic wasps except when larger, more profitable moths are unavailable, consistent with optimal foraging principles (Jones, 1990). However (Swift et al., 1985) found no evidence of prey size selection in foraging pipistrelles.

Invertebrates in Appendix 4 are listed mainly at family level, according to the level of resolution available from dietary analysis. Many insect families comprise species with widely contrasting life-history strategies, habitat requirements and distributions, thus generalisations fit only a proportion of species within any family and not necessarily those species which constitute bat food. Indeed the basic ecology of many invertebrate species remains unknown or only partly described. A characteristic of many insect life-cycles, however, is the utilisation of different habitats in the various life-history stages. Potentially this requires a mosaic of habitats or microclimates to be available within the dispersal range of the different life-history stages. Adult stages of insects preyed on by bats were commonly associated with damp, shady environments such as woodlands, grasslands, freshwater, with live animals or decaying organic matter (Appendix 4). In the larval stage, common habitats of diptera were damp or decaying organic matter or freshwater environments, while Lepidoptera caterpillars were typically associated with a range of foliage types. Dung was a principle habitat for the larvae of many beetles and Diptera. Such habitats are commonly found on agricultural land.

The families preyed on by bats include a substantial number which are mainly diurnal, as well as those with crepuscular or nocturnal activity (Appendix 4). This reflects the gleaning habit of several British bats. Natterer’s bat, an example gleaner, is reported to feed almost entirely on diurnal Diptera harvested from their night resting places (Vaughan, 1997).

3.3 Evidence for past agricultural impact on bat populations

Populations of many bat species are believed to have declined substantially in Britain and across western Europe during the last century, but the lack of attention the group received during this time has led to considerable uncertainty regarding the scale of changes (Moore et al., (submitted); Wickramasinghe et al., In press). Agricultural intensification processes have been proposed as a major driver of declines (Gerell & Lundberg, 1993; Moore et al., (submitted); Wickramasinghe et al., In press). The evidence for this is drawn from a small number of bat studies, bolstered indirectly by a larger body of information relating to the loss or degradation of key habitats (identified in section 3.1) and changes in prey abundance. The mechanisms by which intensive farming is proposed to damage bat populations include reduced survival through loss of roost sites; loss, degradation or reduced accessibility of

11 Project DEFRA title project code foraging areas; depletion of the prey resource and direct or indirect exposure to toxic compounds such as pesticides. Even for historically well-monitored groups such as birds, the simultaneous instigation of many intensification processes has meant that factors contributing to population change cannot usually be separated without experiment (Gillings & Fuller, 1998).

Experimentation and long-term monitoring in relation to agriculture were lacking for bats through most of the twentieth century and the degree to which roosts, foraging habitat and prey abundance limit populations is therefore debated. In a recent attempt to test the hypothesis that agricultural intensification has been a factor in bat declines, (Wickramasinghe et al., In press) compared bat activity on conventional and organic farms, but controlled for landscape and habitat factors. Total bat activity and foraging activity were higher on the organic farms (which were used to simulate less intensive management), but no significant difference in bat species richness was recorded. (Wickramasinghe et al., In press) linked the greater bat activity on organic farms to a greater abundance of aerial insects (Wickramasinghe, unpublished data). Quality of foraging habitat and the abundance of prey has been a consistent theme in studies of bats and aerial-insect eating birds. (Gerell & Lundberg, 1993) attributed pipistrelle declines in industrialized Sweden to deterioration in feeding conditions induced by drainage and water pollution and in similar vein, (Vaughan et al., 1996) noted that British bat declines occurred over the same period as a decline in river water quality. (Arlettaz et al., 2000) proposed competition for a limited food supply as an explanation for the contrasting population dynamics of lesser horseshoe and pipistrelle colonies at a Swiss site. Some of the best evidence for aerial insects as a limiting resource comes from the bird literature, however. In Denmark, abundance of aerial insect-feeding swallows was linked to food supply (Moller, 2001). Swallow food decreased significantly in the absence of cattle, and was reflected in reduced clutch sizes, poorer quality nestlings and fewer adults. Previous evidence for UK swallow declines (Gibbons et al., 1993) is now disputed because new analyses provide no evidence for population change on farmland (Robinson et al., 2003). House martin numbers also appear stable, and population figures for swift are inadequate for analyses of this type (Gibbons et al., 1993). Spotted flycatcher has been in long-term decline since the 1960’s, but this has occurred in both farmland and woodland and the causes are unknown (Gibbons et al., 1993; Freeman & Crick, 2003).

Despite the interest that has been shown in other farmland groups, there is little information concerning long-term trends in invertebrate populations and data are available for relatively few taxa (Sotherton & Self, 2000). The most relevant long-term studies include the Rothamstead suction and moth trap network that has sampled throughout the UK since the mid 1960’s (Woiwod & Harrington, 1993) and the Game Conservancy Trust’s Sussex study (Aebischer, 1991) which has collected data since 1970. The evidence points to a general decline in invertebrates including spiders, leaf beetles and rove beetles (Aebischer & Potts, 1990; Aebischer, 1991), although there is significant variation between regions (Harrington et al., 2003) and invertebrate groups (Aebischer, 1991). (Benton et al., 2002) found temporal correlations between agricultural practice and the abundance of various aerial insects in Scotland and a recent pilot study to expand the analysis of historical trends in aerial insect biomass to four English sites found a significant decline in the Hereford area, but not elsewhere (Harrington et al., 2003). Among the groups of particular importance in bat diet (Appendix 4), moths are reported to have declined widely (Fox, 2001; Harrington et al., 2003) and craneflies (Stubbs, pers. comm.) and dung beetles, (particularly Scarabaeide) (Mann, pers. comm.), have also declined in agricultural landscapes. Although climate and weather effects may have been responsible for some species declines (Conrad et al., 2002), further evidence that agricultural intensification has caused invertebrate population reductions is provided by the many short-term studies that have examined the impacts of typical management practices (reviewed in more detail in section 3.4).

Although there have been recent improvements, detrimental changes in area, character and quality of the main bat foraging and commuting habitats have been recorded since the second world war. Pond numbers fell from 470,000 in 1945 to 243,000 in 1998 and though recent creation programmes may have halted this decline considerable bat and insect habitat would appear to have been lost (Environmental Agency, 2003). Prolonged river water quality declines, primarily associated with agricultural practices, have also undergone a recent slight improvement (references cited in (Vaughan et al., 1996)) but are likely to have influenced local aerial insect populations, and consequently bats. Approximately 250,000 km of hedgerow was removed between 1946 and 1990 (Pollard et al., 1974; Barr et al., 1993). Although (Haines-Young et al., 2000) report indications that the trend has been reversed, the landscape is essentially more open, with less of the linear features known to facilitate the movement of bats between foraging areas such as woodlands. The loss of hedgerow trees during hedge removal, was exacerbated by the impact of Dutch elm disease, reduced bird nesting opportunities (O'Connor & Shrubb, 1986) and probably bat roosts. Changes in woodland area have been more complex, with overall area comprising native and coniferous plantations. The quality of such habitats for bats is likely to have varied dependent on the ratio of deciduous to coniferous woodland, tree age structure, and the abundance of aerial insects. Within the past decade, the area of

12 Project DEFRA title project code woodland on agricultural land has risen by approximately 21% in England and is likely to have instigated further changes in biodiversity.

3.4 Impact of current agricultural practices on bats and their invertebrate prey

3.4.1 Introduction

Following the presentation of general evidence of agricultural impacts on bat populations in the past (section 3.3), this section collates more detailed information on the consequences of individual farming practices. Since very few studies on European bats have investigated the impact of farming at this scale, the main focus is on impacts to bat prey, in the expectation that bats may be affected indirectly, where prey becomes limiting. The invertebrates prioritised in this analysis were Diptera, Lepidoptera and Coleoptera, with an emphasis on those families listed in Appendix 4. Where sources permitted, however, some information on groups such as spiders and Hemiptera (true bugs and plant/leaf hoppers) was also included since these are taken in smaller quantities e.g. (Vaughan, 1997).

Some groups of invertebrates eaten by bats have received less research attention, requiring extrapolation of trends from closely-related groups that are less featured in bat diet. For example, most research on agricultural impacts on Lepidoptera has been directed at butterflies, which are often regarded as useful indicators of environmental change e.g. (Rands & Sotherton, 1986; Davis et al., 1991; Longley & Sotherton, 1997a; Weibull et al., 2000; Saarinen, 2002). Less work has been done on moths in agricultural landscapes and most of this has examined individual pest species, particularly in orchards and forestry (e.g. (Mowat & Clawson, 1988; Cory & Entwistle, 1990; Collier et al., 1996; Mani et al., 1997; Cory et al., 2000; Ioriatti & Angeli, 2002; Degenhardt et al., 2003). Therefore where minimal information on moths existed, impacts on butterflies were reported. Similar issues were encountered with other groups such as beetles, since much of the agricultural beetle literature has examined ground beetles and beneficial predators that are easily sampled with pitfall traps, while other families have received less attention.

Impacts occurring through management of crop and non-crop habitats are considered.

3.4.2 Landscape and cropping pattern

In England agricultural land covers about 70% of the land surface area and is dominated by grasslands (46% of agricultural land) and cereals (30% of agricultural land) (Fig. 1). The remaining area comprises setaside, woodland, horticulture, other land and other arable crops (DEFRA, 2002). The relative proportion of cereals and grassland varies significantly among regions. For example East Anglia comprises 50% cereals and only 14% grassland in distinct contrast to the north-west where 84% is grassland and only 8% cereals. The average size of farm holdings also varies regionally, with holdings larger than 100 ha almost twice as common in the east as in the west (DEFRA, 2002). Larger holdings often arise as part of intensification, through the fusion of smaller units and to optimise the efficient use of machinery, field size in such holdings is often increased by removing boundary features such as hedgerows. The resulting coarse- or fine-grained matrix of arable, grassland and non-crop habitats defines the composition of species assemblages. Non-crop habitats on farmland are usually more species diverse than cropped fields and intensive grassland (Brereton, In press) and may sometimes become islands of species-richness if dispersal across less suitable habitat is limited. In Sweden, (de Jong, 1995) found patterns of bat species richness in forest patches surrounded by agricultural land, consistent with island theory while (Bright, 1993) predicted that several British bats would be vulnerable to habitat fragmentation on the basis of life-history characteristics.

Management practice, vegetation structure and duration to harvest vary among crops and can all affect invertebrate communities (McCracken et al., 2000). Several studies of foraging birds have highlighted differences in invertebrate availability between crop types, e.g. (Green, 1984) found that cereals had higher densities of partridge chick-food insects than sugar beet or carrots, and that winter wheat supported more invertebrates than spring wheat or barley. Crops vary widely in their beetle assemblages and the literature on this group serves to illustrate factors associated with cropping pattern that may be important to invertebrates generally. Patterns in assemblage composition may occur in response to the different spraying regimes associated with particular crops and the amount of ground cover (Hance, 1990). The presence of ground cover promoted carabids (Hummel et al., 2002) and winter wheat can have higher densities of beetles than the early successional stages of set-aside because it provides more cover (Moreby & Aebischer, 1992). Crop type may also influence the relative proportions of day and night active beetles. For instance, the beetle assemblages in maize and ryegrass were dominated by diurnal ground beetle species (Alderweireldt & Desender, 1992).

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The habitat requirements of different invertebrate populations occur at different spatial scales. This results from differences in the mobility of different stages of the lifecycle and differences in the degree to which populations are affected by the timing of certain agricultural operations. At field-scale, ground beetle diversity is highest at the field boundary and decreases towards the field centre (Holland et al., 2001; Haysom et al., In prep). Beetles move from boundary hibernation sites into the field in spring to feed on groups such as aphids. While such affects have been less-studied in other groups they are likely to occur. At the landscape scale, invertebrate diversity is likely to be related to the degree of heterogeneity in crop types and the persistence of certain less mobile species may depend on the presence of less intensively managed crops or non-crop refugia.

Cropping patterns can also affect invertebrate populations in more subtle ways. For instance Diptera such as craneflies are influenced by management that affects vegetation height and density during adult emergence. If oilseed rape preceeds a winter wheat crop, emerging adult craneflies may fail to disperse from under the dense rape canopy and breed locally, causing a build up in population density in the following winter wheat crop (Coll et al., 1993; Coll & Blackshaw, 1996).

There is scope to examine more precisely how the aerial insects eaten by bats and other predators vary with crop type, enabling mitigation priorities for homogenous landscapes with low food-density to be identified.

3.4.3 Farming system

Most of the research that has examined the impact of farming system on biodiversity has compared conventional with organic agriculture. Conventional farming is heavily reliant upon the use of inorganic chemicals for pest control and soil fertility. In contrast, organic farming prohibits the use of synthetic fertilisers and pesticides, relying instead on more traditional mechanical techniques. It is difficult to make general statements about the impacts that different farming systems are likely to have on bat-food insects. The wide use of synthetic pesticides in conventional farming is likely to have had the greatest impact on invertebrates and is addressed separately (see 3.4.4). Apart from the prohibition of synthetic pesticides organic farming is generally less intensive and relies on more traditional techniques. These include crop rotations, crop diversification, green manures/leys, farmyard manure (FYM) and composts to maintain soil fertility, and the enhancement of natural enemy populations (e.g. predatory beetles, bugs and parasitoid wasps), particularly through sympathetic boundary management. These elements are likely to have fewer negative consequences for invertebrates and some practices may even be beneficial. However, some organic farms can be fairly intensive. For example nearly half the hedgerows removed by farmers during the mid 1990's were removed by organic farmers (Stoate et al., 2001). In addition, organic farming relies upon mechanical weed control including soil cultivation such as ploughing which can be very destructive (Stoate et al., 2001), particularly to soil dwelling invertebrates (Frouz, 1999).

Results of studies comparing invertebrate biodiversity in organic and conventional farming systems are not conclusive. Organic farming has been reported to benefit ground beetles (Shah et al., 2003), spiders (Marc et al., 1999) and butterflies (Feber et al., 1997), although (Weibull et al., 2000) found no significant differences between butterfly populations on organic and conventional farms. Other invertebrate orders such as staphylinid beetles (Shah et al., 2003) have been found to be more abundant in conventional fields. Possible explanations for this are that staphylinids, a group of highly mobile beetles, may avoid pesticide applications more easily than other less mobile groups, that the higher density crop in conventionally managed fields creates a more humid microclimate which suits this group, or that some of the species with which staphylinids compete are less abundant in conventionally managed fields. Some of the more recent literature attributes many of these patterns to landscape heterogeneity rather than management differences per se. For instance, (Weibull et al., 2000)) found that invertebrate abundance was positively correlated with heterogeneity within the farm and that species diversity was positively correlated with the diversity of field types in the surrounding landscape.

(Wickramasinghe et al., In press) found that bat activity was 61% higher, and foraging activity 84% higher on organic than conventional farms. (Wickramasinghe et al., In press) suggested that the habitats found on the organic farms were of higher quality in terms of habitat structure and condition (due to the lack of agrochemicals), supported higher overall insect abundance and more of the key insect families important to bats as food. Preliminary results of an ongoing study (Mathews, pers. comm) support these findings, with significantly higher bat activity recorded on transects through organic, than conventional farms.

3.4.4 Impact of agrochemicals on biodiversity

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A wide range of different pesticides (principally herbicides, insecticides and fungicides) are used on crops grown in the UK and the application rates vary between crops. For example herbicide applications can vary from one application annum-1 on rye and triticale (Garthwaite & Thomas, 2000) to six applications annum -1 on onions and leeks (Garthwaite et al., 1999). Pesticide usage in grassland occurs, but levels are insignificant compared to the amounts applied to arable crops (Vickery et al., 2001). Pesticides may reduce invertebrate abundance through direct toxicity, but also indirectly by restricting food supply or altering habitat.

The impact of pesticides on different invertebrate groups is highly dependent upon the timing of application (Ewald & Aebischer, 1999). For example, autumn applications tend to drift more because crop and marginal vegetation heights are lower (Longley & Sotherton, 1997b) and may therefore contact invertebrates over a larger area. During recent decades herbicide use has increased and switched gradually from spring/summer applications to autumn/winter applications, reflecting the dramatic trend towards autumn sown crops over the last 30 years. Although the use of other pesticides has also increased, the timing of application still varies greatly between crops (Ewald, 2000).

Most of the research that has examined the impact of pesticides on invertebrates has focused on insecticides and herbicides. Although fungicide use is still increasing relatively little research has considered its impact. Existing sources indicate that some fungicides reduce the density of several invertebrate groups including staphylinids, carabids (Sotherton et al., 1987), aquatic organisms (de Snoo, 1999) and spiders (Ewald & Aebischer, 1999). Mechanisms are likely to include direct toxicity and a reduced food supply since moulds are important in the diet of many detritivorous species.

Insecticides are reported to reduce the abundance and species richness of some dipteran families, particularly those with soil dwelling larvae (Frouz, 1999), in crops (Vickerman & Sunderland, 1977; Fischer & Chambon, 1987), water (Ward et al., 1995; Woin, 1998) and grassland habitats (Linzell & Madge, 1986). Some families of flies such as hoverflies are very sensitive to insecticides (Fischer & Chambon, 1987; Theiling & Croft, 1988), whilst others such as nematocera (e.g. midges and craneflies), have shown a mixture of responses depending upon the habitat and the type of application. For example, chironomid larvae in ponds that received a single pesticide application (Woin, 1998), or in wheat fields which received several aphicide applications (Fischer & Chambon, 1987), showed little response, in contrast to the situation in streams where low level continuous applications greatly reduced populations (Ward et al., 1995).

Few papers refer to the impacts of herbicides on Diptera (Dewey, 1986; Cowgill et al., 1993; Moreby et al., 1994; Moreby & Southway, 1999). Herbicides appear to depress Diptera abundance indirectly, both on land (Cowgill et al., 1993; Moreby & Southway, 1999) and in ponds (Dewey, 1986), by reducing the availability of plants utilised as food or microhabitat. For example hoverflies were more abundant in headlands that were not sprayed by herbicides (Cowgill et al., 1993). This is likely to have been an indirect effect delivered through a reduction of the available pollen sources that are known to be a factor limiting egg maturation in females, especially in agricultural landscapes (references in (Wratten et al., 2003)).

Non-pest moth assemblages have been poorly recorded on agricultural land and the only evidence available suggests that pesticides have a negative impact, at least on larvae (Ewald & Aebischer, 1999). However, potential agrochemical impacts on moths may be extrapolated from the butterfly literature. Pesticide use is considered to have had a major impact on farmland butterfly populations (Rands & Sotherton, 1986; Longley & Sotherton, 1997a; de Snoo, 1999; Ewald & Aebischer, 1999; Brereton, In press). As many butterflies in agricultural landscapes are associated with non-crop boundary vegetation (Brereton, In press) much research has addressed the potential impact of spray drift into field edges and hedgerows. The evidence suggests that butterflies are at risk from spray drift in conventional fields (Longley et al., 1997; Longley & Sotherton, 1997b) and that a margin of unsprayed crop acting as a buffer zone reduces the risk from spray drift significantly (Longley et al., 1997; Longley & Sotherton, 1997b; de Snoo, 1999). However, there is still some debate over the width of buffer required. Depending on the intended function, agri-environment schemes prescribe various widths of unsprayed boundary. Some evidence suggests that a 6m buffer is insufficient to eliminate risk, especially in strong winds (Longley & Sotherton, 1997b), whilst other studies imply that a 6m buffer conveys no more protection than a 3m buffer (de Snoo, 1999). An explanation for this inconsistency may be that the nature of the crop, especially density and height, influences the amount of spray reaching field boundaries (Longley et al., 1997).

Herbicides reduce butterfly abundance indirectly when they cause the loss of larval food-plants and nectar or pollen sources for adults, particularly in boundary vegetation (Clausen et al., 2001). There is some evidence that population change in some butterfly species is linked to changes in plant populations on agricultural land (Smart et

15 Project DEFRA title project code al., 2000). However, the only clear associations are for species of butterfly that are increasing in response to improved availability of larval food plants such as nettles, which are encouraged by fertilisers (Smart et al., 2000).

Pesticides are generally reported to have a negative impact on beetles (Vickerman & Sunderland, 1977; Booij & Noorlander, 1988; Clements et al., 1988), including their larval stages (Vickerman & Sunderland, 1977; Linzell & Madge, 1986). In most cases pesticides are reported to cause population declines over short periods (Edwards & Thompson, 1975; Vickerman & Sunderland, 1977; Booij & Noorlander, 1988; Cilgi & Frampton, 1994), but there may also be longer term implications. For example, long term studies have shown that large-scale increased usage of insecticides can have a severe impact on ground beetle abundance, with some species disappearing from intensively managed fields (Basedow, 1990). Pesticide toxicity can act on beetles during application or afterwards through residual exposure from contaminated vegetation and soil. Residual exposure is deemed to be the most important pathway for ground beetles (Thacker & Hickman, 1990) although indirect effects through changes in prey population are also important (Basedow, 1990). Herbicides are known to impact on beetles through changes in plant cover, particularly important for phytophagous beetles such as weevils (Curculionidae) and leaf beetles (Chrysomelidae) (Moreby & Southway, 1999). There is often variation between families (Moreby et al., 1994) and species (Asteraki et al., 1992; Krooss & Schaefer, 1998) with some studies reporting no overall impacts (Fischer & Chambon, 1987; Hummel et al., 2002). The impacts on different sexes may also differ (Chiverton & Sotherton, 1991).

The timing of pesticide application, emergence of beetles, their diurnal activity patterns and size can have an influence on the magnitude of the effects of pesticides e.g. (Critchley, 1972; Gregoire-Wibo & Hoecke, 1979). Species overwintering in fields are more likely to be exposed to pesticides when little or no vegetation cover is available (Cilgi & Frampton, 1994). For ground beetles it has been shown that the effects of pesticides in the centre of the field are more severe and occur faster than those at the edges of the sprayed area (Duffield & Baker, 1990).

Insecticide application has generally been found to reduce spider density (Vickerman & Sunderland, 1977; Fischer & Chambon, 1987; Krause, 1987; Cilgi & Frampton, 1994; Moreby et al., 1994; Thomas & Jepson, 1997; Marc et al., 1999; Hummel et al., 2002) and diversity (Pekar, 1999) in both crop and non-crop habitats. Most of the impacts act indirectly by altering the abundance and composition of prey populations and spider populations usually recover within a few months (Cilgi & Frampton, 1994; Thomas & Jepson, 1997; Marc et al., 1999). Timing of application seems to be important, with summer applications having less impact (Cilgi & Frampton, 1994). Much of the literature focuses on ground active spiders such as linyphiids and lycosids that are less likely to appear in the diet of bats. However, those studies that have looked at vegetation dwelling spiders, which are more likely to be gleaned by bats, showed similar trends (Krause, 1987; Marc et al., 1999).

Herbicides also reduce spider densities (Krause, 1987; Moreby et al., 1994; Haughton et al., 1999a; Haughton et al., 1999b; Moreby & Southway, 1999; Haughton et al., 2001; Bell et al., 2002) via changes in vegetation composition and structure (Krause, 1987). As with insecticides, spider populations can recover after single applications with often no significant impacts detectable by the next season (1 year: (Bell et al., 2002); 16 months: (Haughton et al., 2001)). However, most research has been done on ground active spiders and those reliant on the presence of vegetation for building webs may be slower to recover. On most farms, herbicides are applied several times a year, depending on the crop type (Garthwaite et al., 1999; Garthwaite & Thomas, 2000) and this has the potential to inhibit the recovery of spider populations over much longer periods of time.

In addition to potential impacts on bats via reduced food supplies, some authors consider the potential susceptibility of bats to pesticide poisoning, though direct contact with sprayed vegetation or through eating sprayed insects or insects that have eaten contaminated food e.g. (Duvergé & Jones, 2003). The extent to which this actually occurs under field conditions is poorly documented however and Cantwell (pers. comm.) reports that the number of wildlife incidents associated with pesticide use in general have fallen since the early 1990’s. Studies from the 1970’s and mid 1980’s cited in (Duvergé & Jones, 2003) documented instances of pesticide residue accumulation in bats. Another study that collected data during the 1980’s reported higher concentrations of organochlorine residues in bats from an industrialized area of Sweden, than in a rural area (Gerell & Lundberg, 1993). While residues occurred at non-lethal levels, potential alterations to fat metabolism leading to premature fat depletion during hibernation were discussed.

3.4.5 Avermectins

Invertebrates associated with cattle dung feature in the diet of several bats. Pasture invertebrate assemblages are potentially threatened by modern livestock endo-parasite control practices. The best studied group of endo-parasitic

16 Project DEFRA title project code treatments, in terms of potential environmental impacts, is the avermectins, a group of synthetic pyrethroids which have been widely used for the last twenty years. Whilst these chemicals are effective at endo-parasite control, they do not decompose well and can remain active in dung at least 5 weeks after treatment (Spratt, 1997). Avermectins and similar compounds are administered to cattle topically (pour on), by injection or as a slow release bolus. Pour on or injection can be applied as a single dose or three doses 3, 8 and 13 weeks after animals are turned out (McCracken, 1995). Bolus application results in a slow release of avermectins over 135 days. For all treatments whilst the avermectin is active in the animal it will be present in the dung. The extent to which avermectin persistence in dung represents a threat to non-target organisms has been widely debated.

A literature review by (Strong, 1992; Strong, 1993) suggested that avermectins were harmful to non-target dung invertebrates, especially higher flies, at concentrations found in dung several weeks after application. Dung flies are particularly sensitive to avermectins with severe mortality in both larval and adult stages reported up to 30 days after application (Sommer et al., 1992; Strong, 1992; Strong, 1993; Lumaret & Kadiri, 1998). The impact on invertebrates varies with treatment method. Application by injection is reported to have longer lasting impacts on dung fauna than application by pour on (Sommer et al., 1992) and dung from bolus treated cattle is reported to have particularly high toxicity (Gover & Strong, 1996). Primitive flies such as Nematocera are also killed by avermectin contaminated dung but the effects occur over shorter periods (Madsen et al., 1990; McCracken & Foster, 1993). Beetles, especially dung beetle larvae, also suffer mortality in the dung of avermectin treated animals, but the effects are short-lived, usually over several days (Madsen et al., 1990; McCracken & Foster, 1993; Strong, 1993).

In addition to direct mortality, avermectins cause non-lethal effects such as reduced invertebrate fecundity which are often overlooked. Non-lethal effects would appear to act over longer periods (Strong, 1992; Strong, 1993; Gover & Strong, 1996), and continued exposure to avermectins could depress sensitive populations of dung beetles such as Aphodius sp (Mann, pers. comm.).

The key issues with avermectin use are the type and timing of application as many are applied during the peak activity of invertebrates and feeding by birds and mammals (McCracken, 1993). Bolus or repeated injections over the summer are likely to have greatest impact (McCracken, 1995). However, although there is strong evidence that avermectins can be detrimental to certain invertebrate populations the use of endo-parasitic products is crucial to livestock farming (McCracken, 1995). Furthermore, some authors, reported in (Vickery et al., 2001), have suggested that because there is sufficient uncontaminated dung at the landscape level the impact of avermectins and associated products on populations of dung fauna is likely to be quite limited, although local impacts could be severe. The widespread reduction in quality of dung in improved pastures for diptera (Stubbs, pers. comm.) and dung beetles (Mann, pers. comm.) may exacerbate the impact of avermectins on invertebrate populations at larger scales. Advances in veterinary science may provide a solution as alternative compounds, which combine the benefits of avermectins with insignificant impacts on dung fauna, become more available (Strong & Wall, 1994; Strong et al., 1996).

The results of a study that examined activity of different bat species in relation to avermectin use were inconclusive overall (Duvergé, pers. comm.). However, since juvenile greater horseshoe bats target large, slow-moving beetles associated with cowpats, proposals to limit the use of avermectins near maternity colonies were made as a precautionary principle, and have been widely instigated (Duvergé, pers. comm).

3.4.6 Fertiliser practices

Synthetic fertiliser application on tillage crops has seen a slight downward trend since a thirty-year peak in the mid 1980's of on average 157 Kg Nitrogen ha-1 year-1 falling off to an average of 149 Kg Nitrogen ha-1 year-1 applied in 2000. Nitrogen inputs on grassland have also declined since the mid 1980's when in 1984 the average application peaked at 131 Kg Nitrogen ha-1 year-1, falling to 99 Kg Nitrogen ha-1 year-1 in 2000 (Chalmers et al., 2001).

Increased use of inorganic fertilisers generally occurs at the same time as intensification of other management practices such as grazing and mowing, so it is often difficult to separate out direct fertiliser effects in monitoring studies. A recent review (Vickery et al., 2001) highlighted a range of invertebrate responses to fertiliser applications. Leatherjackets (cranefly larvae) appear indifferent to artificial fertiliser levels, whereas plant feeding true bugs, plant hoppers and leaf hoppers respond positively to increased nitrogen (Whittaker, 1976; Andrzejewska, 1976; Sedlacek et al. 1988; Morris, 1992) and some flies and beetles suffer population reductions under high nitrogen regimes (Vickery et al., 2001). High nitrogen can be detrimental to larger insects such as the larger species of ground beetle, especially in conjunction with intensive grazing and/or mowing (Blake et al., 1994; Fuller, 2003).

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Impact on phytophagous species can be linked to the reduction in plant species richness that occurs in high fertility soils e.g. (Fuller, 2003).

The use of organic fertiliser such as FYM and slurry also elicits complex responses from the wide variety of invertebrates found in grasslands (Curry, 1987). Organic fertilisers generally benefit dung fauna including groups of flies (e.g. Bibionids and Tipulids) and beetles important to bats. However, this depends on the source of the dung, with cattle manures being preferable to pig manure, and on the level of use. Excessive use of slurries can impact on some groups through oxygen depletion and toxicity of chemicals in the slurry. Several bat species are reported to feed on insects at manure piles e.g. (McAney & Fairley, 1988). Correct management of all types of fertilizer on farmland to minimize nitrogenous runoff are important , since bat foraging activity may be decreased significantly in polluted water (Vaughan et al., 1996).

3.4.7 Cutting and grazing

Grassland managements such as grazing and cutting for hay or silage alter the structure and composition of vegetation and can have major impacts on invertebrate abundance and species composition (Curry, 1987; Gibson et al., 1992; Dennis et al., 1998; Morris, 2000). Generally, undisturbed tall grassland with a complex structure supports a greater abundance and diversity of invertebrates than short uniform grassland (Curry, 1987). The frequency and timing of cutting and grazing are influential (Morris, 2000) as the consistent repetition of management from year to year can select for different invertebrate assemblages if, for example, mowing removes vegetation and animals before they are able to breed.

The impact of cutting and grazing on Diptera is difficult to assess as many studies have been at order level and consequently combine resident field species and visitors. There is some evidence to suggest that higher densities of flies occur in heavily grazed fields (Gibson et al., 1992), but it is likely that these are drawn by the presence of livestock and dung. A review by (Gerstmeir & Lang, 1996) indicated that cutting twice per year was detrimental to hoverflies but for other families responses were mixed. Vegetation structure and density also influences Diptera abundance. The succession process that occurs on set-aside land with bare ground being replaced by dense vegetation can lead to reduced population densities of chironomids (Frouz, 1994).

The timing and frequency of grassland management seems to be critical for beetles. For example, (Fuller, 2003) found that several silage cuts per year had a greater impact on beetles, particularly phytophagous species, than one late hay cut. An extensive review by (Gerstmeir & Lang, 1996) on the effects of mowing on arthropods clearly highlights the difficulty of generalising about a large order such as beetles. While the overall abundance of beetles was not greatly affected by cutting, both positive and negative responses were recorded within the same families. (Gibson et al., 1992) suggests that short-term spring grazing benefits some beetles. Even when conservation is the primary objective of management, conflict can occur with practices that benefit other taxa. For example, e.g. on nature reserves managed for botanical diversity, cattle are often removed in the summer. This removes a vital food resource (dung) at a crucial time for dung beetles although species with sufficient mobility may relocate to improved pasture. There is currently speculation that the nitrogen rich diet of cattle feeding on improved pasture results in a more liquid dung which is a poorer resource for most dung beetles, especially Scarabaeids (Mann, 2003).

Many butterflies are associated with grassland habitats and the loss of semi-natural grasslands is often cited as a major cause of species declines (Brereton, In press). Many species are sensitive to grassland management. Responses to grazing are complex as this may affect several life history stages i.e. it can remove both larval food resources and nectar sources for adults. Both over- and under- grazing are reported to have affected certain species. Heavy grazing that produces a uniform short sward is generally detrimental to butterflies (Saarinen, 2002) whereas light grazing can be beneficial if it results in a mosaic of different vegetation heights and densities (Clausen et al., 2001). Individual species requirements may be very specific and in the case of chalk grassland butterflies some species require heavy grazing and others light or no grazing (Thomas, 1987). Cutting acts in a similar way to grazing but the uniform nature of mowing can lead to greater impacts because all the vegetation is reduced at one time. The timing of cutting is crucial to butterflies. Authors reviewed in (Gerstmeir & Lang, 1996) reported that mowing was detrimental to blue butterflies (Lycaenidae) but had no effect on browns (Satyridae). Another agricultural grassland practice, reseeding with species-poor cultivated ley mixtures, can reduce Lepidoptera diversity, because of the reduced diversity of larval food-plants.

Grassland management such as grazing (Gibson et al., 1992), and silage cutting (Thomas & Jepson, 1997) can seriously reduce spider abundance with recovery often taking several months (Thomas & Jepson, 1997). Intensive

18 Project DEFRA title project code grazing can lead to virtual extinction (Thomas & Jepson, 1997). Increasing the structural complexity of vegetation, e.g. by planting grasses in orchards, can lead to increased abundance of arboreal spiders (Pekar, 1999).

Many true bugs (Hemiptera) are closely associated with grasslands and several authors have studied the effects of management on this order, particularly true bugs and leaf and plant hoppers. Tall swards have been identified as supporting more species and larger populations than those reduced in height by cutting (Morris, 1981a) and grazing (Dennis et al., 1998). Grazing by sheep can be particularly damaging with up to 10-fold reduction in the abundance of common Heteroptera in sheep grazed plots (Gibson et al., 1992). Less work has been done on grazing by cattle although recently (Fuller, 2003) suggested that high intensity cattle grazing, and silage cutting, reduces numbers of plant and leaf hoppers (Auchenorrhyncha). However, intensive management of this kind was associated with high nitrogen inputs which other authors have suggested increase hopper numbers (Whittaker, 1976; Andrzejewska, 1976; Sedlacek et al. 1988; Morris, 1992). Numbers of heteroptera were too low to analyse but appeared to occur less frequently in intensively managed pastures. Mowing has also been shown to impact on both Heteroptera and Auchenorrhyncha (Gerstmeir & Lang, 1996) but responses vary between species and timing of the cut (Morris, 1981a; Morris, 1981b). Generally cutting reduces hemipteran biomass particularly if the cut occurs in July. Effects persist for some time in species which overwinter as adults (Morris, 1981a).

3.4.8 Drainage

The most important agricultural intensification impact on Diptera was probably the widespread installation of field under-drainage during the 1970's and 1980's (Stubbs, 2001). This occurred during the period of substantial modification to river catchments in southern Britain that also resulted in the loss of important dipteran habitats such as sand and shingle shoals on river banks (Stubbs, 2001). Another important change for diptera has been the shift from a 3-4 year rotational hand ditch clearance system to regular large-scale mechanical clearance leaving deep, steep sided ditches. This has resulted in large networks of ditches which are unsuitable for the majority of species which require bankside sediments and vegetation (Stubbs, 2001). Drainage is also reported to have affected the abundance of moths (Fox, 2001) and Hemiptera (Kirby et al., 2001).

3.4.9 Soil cultivation

Tillage, particularly ploughing, can cause high mortality in populations of soil dwelling Diptera larvae (Frouz, 1999; Frouz & Paoletti, 2000).

Soil cultivation tends to reduce populations of beetle larvae (Krooss & Schaefer, 1998) and can influence community structure (Purvis et al., 2001) by selecting against species with long larval stages. Since species with longer lived larvae are often larger bodied, this can result in intensively managed habitats being characterised by smaller beetles than extensively managed habitats (Blake et al., 1994; Fuller, 2003). Effects vary with time of disturbance and between families and species of beetle e.g. (Hummel et al., 2002; Purvis & Fadl, 2002; Holland & Reynolds, 2003). Several of the beetle groups with long-lived larvae (e.g. cockchafers and dung beetles) feature prominently in the diet of bats such as serotine and greater horseshoe. For these species, spatial heterogeneity of cultivation management is desirable at farm and landscape level to encourage continuous food supplies. In contrast to large-scale cultivation, mechanical weed control (used widely in organic systems) has minmal impact on ground beetles and rove beetles (Krooss & Schaefer, 1998). moths-ploughing (Fox, 2001)

Cultivation such as ploughing has a strong negative impact on spiders (Marc et al., 1999) with autumn cultivation being damaging (Thomas & Jepson, 1997). Ploughing is also reported detrimental to moths (Fox, 2001) and Hemiptera (Kirby et al., 2001).

3.4.10 Management of hedgerows

Hedgerows provide a broad range of resouces for invertebrates which may use them as courtship and breeding sites, feeding sites, as refuges from predators, hibernation sites or as shelter. Hedges are particularly important for butterflies (Clausen et al., 2001; Brereton, In press) and moths (Bowden & Dean, 1977; Luff & Woiwod, 1993) in agricultural landscapes. Hedge height, width, density, the presence of trees and the composition of the ground-layer vegetation strongly influence the associated invertebrate assemblage. For example, some butterfly species prefer unkempt hedges with uncultivated margins (Brereton, In press). Management that affects hedge structure has complex impacts on invertebrates (Maudsley, 2000). Generally cutting is detrimental, especially to mobile groups such as diptera, and the frequency and timing of cuts is important.

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Hedge height and permeability affect the aerodynamic properties of hedges and can strongly influence the numbers of passing insects that get caught in eddies on the leeward side, so that large numbers may accumulate along tall dense hedges (Lewis & Stephenson, 1966; Lewis, 1969; Lewis & Dibley, 1970; Bowden & Dean, 1977). The structure of hedges can also influence the Lepidoptera fauna (Brereton, In press). However, the vast majority (over 90%) of insects found near hedges do not originate in the hedge but come from other habitats brought in on the wind (Lewis, 1969).

Hedgerows both facilitate and restrict invertebrate movements and flight activity (Frouz & Paoletti, 2000). For example, butterflies disperse along linear features such as hedgerows, but may also perceive them as barriers to movement (Dover & Fry, 2001) and even normal hedgerows may restrict movements of hoverflies between fields (Wratten et al., 2003). Hedgerows can also restrict the movement of beetles such as carabids causing aggregations around field boundaries (Mauremootoo et al., 1995; Holland et al., 2001). Aspects of hedgerow structure also appear to facilitate use by bats. (Wickramasinghe et al., In press) reported a significant correlation between the number of feeding buzzes and hedgerow height. (Duvergé & Jones, 2003) reported the use of thick, well developed boundaries by greater horseshoes.

4. Provision for bats in agri-environment schemes

Appendices 4 and 5 summarise the range of prescriptions available in various agri-environment schemes currently in operation throughout the UK. In England, agri-environment schemes are currently being reviewed and revised with the intention of encouraging wider implementation of conservation measures on farmland in the future. The tables summarise general actions available for habitats particularly associated with roosting or foraging bats (see section 3.1). Although it is known that schemes tend to be habitat rather than species focused, it is notable that very few of the options are described as important to mammals in scheme literature. Only the Scottish Rural Stewardship Scheme water margin management options, and the Northern Ireland Environmentally Sensitive Area Scheme (Appendix 5) specifically refer to bats in the general scheme booklet. In contrast many of the prescriptions list birds among the benefiting species and several groups such as waders have options specifically targetted toward their conservation.

Since prescriptions are succinct, allowing advisors some flexibility to tailor recommendations to local circumstances, land managers with a particular interest in bat conservation should refer to a more detailed text e.g. (Entwistle et al., 2001).

5. Recommendations

5.1 Opportunities for future research

 Improved resolution of dietary information would enable more precise management recommendations to enhance the populations of prey groups.

 Greater integration of research at water edges to include impacts of management on biodiversity associated both within the watercourse and with surrounding vegetation.

 Examination of how habitat use, colony size and distance travelled to foraging sites varies among bat colonies in different landscapes, particularly along an arable predominant – mixed farming - grassland predominant gradient.

 Greater examination of population status, changes and pressures acting on bats in predominantly arable landscapes.

 Examination of large-scale monitoring data to determine how measures of bat-activity vary with fragmentation of key habitats. Use of modelling and economic approaches to consider how favourable habitat mosaics could be implemented at landscape scale.

 Greater use of radio-tracking to examine the use of agricultural landscapes by different species of bats and to test impact of agri-environment scheme prescriptions.

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 There is a paucity of studies addressing population changes in Diptera, or quantitative information on variation in abundance among habitat types. Further research, including manipulative experiments, examining the impact of management on diptera populations would benefit the conservation of groups that predate them e.g. bats, hirundines. Research would need to address impacts at key lifecycle stages.

 The impact of habitat management on the abundance and species composition of moth assemblages is under-researched. Co-ordinated work on trends in moth populations (both night-flying and day-flying species) in association with direct work on management impact, would be of benefit.

 Broader research on the impact of changes in livestock husbandry and management on the invertebrate assemblages associated with dung. Consideration should be given to dung abundance, continuity of the dung resource, dung quality and species composition of the associated fauna.

5.2 Bat conservationists / SNCO’s

 Evaluate mechanisms by which records of species distributions, roost locations etc could be improved by increasing record coverage and resolution.

 Evaluate pathways to facilitate greater availability of fine-scale species distribution and roost data to enable greater provision for bats in individual agri-environment agreements.

 Promote bat conservation to farmers and land-managers, land-agents, agricultural colleges and agri- environment advisors.

5.3 Land managers

 Report location of bat roosts and observations of flying bats to agri-environment advisors when seeking conservation advice.

 Trees may harbour bat roosts. Avoid removing standing deadwood and conduct tree-work according to Arboricultural Association guidance note.

 Traditional buildings such as barns may be used by bats throughout the year. Seek advice prior to timber treatment and building work to ensure due consideration for bats.

5.4 Agri-environment professionals

 When considering the existing and potential wildlife value of farmland note that land with water, woodland and connective linear features is used by bats. Create and restore these features where possible.

 Consider the presence of bats in features that are managed primarily for their historical /archaelogical features and the potential value of mines and underground sites as hibernacula. Ensure any measures to seal or gate entrances to such sites consider their potential use by bats, so that sites remain available.

5.5 Policymakers

Encourage the use of consortia applications, or target applications from areas in bat "hotspots" to provide greater continuity of habitat.

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Acknowledgements We are grateful to the many conservationists, researchers, FWAG and SAC advisors, RDS ecologists and biodiversity co-ordinators who supported the authors by giving generously of their time in useful discussions, or making unpublished data available. Particular thanks are due to Gareth Jones, Laurent Duvergé, Ian Davidson- Watts, Liat Wickramasinghe, Nancy Vaughan, Fiona Mathews, Niall Moore, Patty Briggs, Stephanie Wray, James Aegerter, Andy Hoodless, Alan Stubbs, Darren Mann, Tom Brereton, Tim Thom, Mark Stevenson, Chris Kaighin, Robert Goodison, Iain Diack, Adam Kwolek, Ian Johnson, Phil Tolerton, Siobhan Murphy, Brian Cantwell, Nick Thomas, Bruce Philp, Davy McCracken, Anne Heeley, Alison McKnight, Tommy Loudon, George Dodds, Seamus Eaves, Martin Phillips, Barney Parker, Mike Williams, Ros Willder, Mathew O’Brien, Elizabeth Ranelagh, Roland Stonex, Mary Combe, Paul Holmes-Ling, Alex Long, Richard Roberts, Helen Bibby, Paul Chapman, Sandra Stewart and Alasdair Smithson.

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Thacker, J. R. M. & Hickman, J. M. (1990). Techniques for investigating the routes of exposure of carabid beetles to pesticides. In The role of ground beetles in ecological and environmental studies. (ed. N. E. Stork), pp. 105-113. Intercept, Andover. Theiling, K. M. & Croft, B. A. (1988). Pesticide side-effects on arthropod natural enemies: a database summary. Agriculture, Ecosystems and Environment 21, 191-218. Thomas, C. F. G. & Jepson, P. C. (1997). Field-scale effects of farming practices on linyphiid spider populations in grass and cereals. Entomologia Experimentalis Et Applicata 84, 59-69. Thomas, J. A. (1987). The conservation of Adonis Blue and Lulworth Skipper butterflies - two sides of the same coin. In Calcareous Grasslands - Ecology and Management (ed. S. H. Hillier, D. W. H. Walton and D. A. Wells), pp. 112-117. Bluntisham Books, University of Sheffield. Vaughan, N. (1997). The diets of British bats. Mammal Review 27, 77-94. Vaughan, N., Jones, G. & Harris, S. (1996). Effects of sewage effluent on the activity of bats (Chiroptera: Vespertilionidae) foraging along rivers. Biological Conservation 78, 337-343. Vaughan, N., Jones, G. & Harris, S. (1997). Habitat use by bats (Chiroptera) assessed by means of a broad- band acoustic method. Journal of Applied Ecology 34, 716-730. Verboom, B. & Huitema, H. (1997). The importance of linear landscape elements for the pipistrelle Pipistrellus pipistrellus and the serotine bat Eptesicus serotinus. Landscape Ecology 12, 117-125. Verboom, B. & Spoelstra, K. (1999). Effects of food abundance and wind on the use of tree lines by an insectivorous bat, Pipistrellus pipistrellus. Canadian Journal of Zoology-Revue Canadienne De Zoologie 77, 1393- 1401. Vickerman, G. P. & Sunderland, K. D. (1977). Some effects of Dimethoate on arthropods in winter wheat. Journal of Applied Ecology 14, 767-777. Vickery, J. A., Tallowin, J. R., Feber, R. E., Asteraki, E. J., Atkinson, P. W. & Brown, V. K. (2001). The management of lowland neutral grasslands in Britain: effects of agricultural practices on birds and their food resources. Journal of Applied Ecology 38, 647-664. Walsh, A. L. & Harris, S. (1996a). Foraging habitat preferences of vespertilionid bats in Britain. Journal of Applied Ecology 33, 508-518. Walsh, A. L. & Harris, S. (1996b). Factors determining the abundance of vespertilionid bats in Britain: Geographical, land class and local habitat relationships. Journal of Applied Ecology 33, 519-529. Ward, S., Arthington, A. H. & Pusey, B. J. (1995). The effects of a chronic application of Chlorpyrifos on the macroinvertebrate fauna in an outdoor artificial stream system: species responses. Ecotoxicology and Environmental Safety 30, 2-23. Warren, R. D., Waters, D. A., Altringham, J. D. & Bullock, D. J. (2000). The distribution of Daubenton's bats (Myotis daubentonii) and pipistrelle bats (Pipistrellus pipistrellus) (Vespertilionidae) in relation to small-scale variation in riverine habitat. Biological Conservation 92, 85-91. Waters, D., Jones, G. & Furlong, M. (1999). Foraging ecology of Leisler’s bat (Nyctalus leisleri) at two sites in southern Britain. Journal of Zoology 249, 173-180. Weibull, A.-C., Bengtsson, J. & Nohlgren, E. (2000). Diversity of butterflies in the agricultural landscape: the role of farming system and landscape heterogeneity. Ecography 23, 743-750. Wickramasinghe, L. P., Harris, S., Jones, G. & Vaughan, N. (In press). Bat activity and species richness on organic and conventional farms: impact of agricultural intensification. Journal of Applied Ecology . Woin, P. (1998). Short- and Long-Term Effects of the Pyrethroid Insecticide Fenvalerate on an Invertebrate Pond Community. Ecotoxicology and Environmental Safety 41, 137-146. Woiwod, I. P. & Harrington, R. (1993). Flying in the face of change: Rothamstead insect survey. In Long-term experiments in agricultural and ecological sciences. (ed. R. A. Leigh and A. E. Johnston), pp. 321-342. CAB International, Rothamstead. Wratten, S. D., Bowie, M. H., Hickman, J. M., Evans, A. E., R.J., S. & Tyliankis, J. M. (2003). Field boundaries as barriers to movement of hover flies (Ditpera: Syrphidae) in cultivated land. Oecologia 134, 605-611.

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APPENDICES

Appendix 1. Habitats identified in (JNCC, 2001). as priority habitats for the six bat BAP species. Habitats closely associated with agriculture are shown in bold. Greater mouse-ear bat is believed extinct in the UK and is distinguished from the five extant species by the use of parentheses. Species BAP Priority habitats BAP Broad habitats  Cereal field margin  Boundary & linear features  Ancient &/or species rich hedgerows  Arable and horticulture  Aquifer fed naturally fluctuating water  Rivers & streams bodies  Calcareous grassland  Coastal floodplain & grazing marsh  Coniferous woodland  Eutrophic standing waters  Broad-leaved & mixed yew woodland Pipistrelle  Lowland beech & yew woodland  Dwarf shrub heath Pipistrellus pipistrellus  Lowland calcareous grassland  Fen, marsh & swamp  Lowland heath  Neutral grassland  Lowland meadow  Standing open water & canals  Lowland wood-pasture & parkland  Improved grassland  Mesotrophic lakes  Native pine woodlands  Wet woodlands  Cereal field margin  Boundary & linear features Lesser horseshoe  Ancient &/or species rich hedgerows  Arable and horticulture Rhinolophus  Chalk stream  Rivers & streams hipposideros  Lowland beech & yew woodland  Broad-leaved & mixed yew woodland  Wet woodland  Ancient &/or species rich hedgerows  Boundary & linear features  Lowland calcareous grassland  Calcareous grassland Greater horseshoe  Lowland meadow  Broad-leaved & mixed yew woodland  Wet woodland  Improved grassland R. ferrumequinum  Maritime cliffs & slopes  Neutral grassland  Lowland wood-pasture & parkland

 Coastal floodplain & grazing marsh  Calcareous grassland  Lowland calcareous grassland  Broad-leaved & mixed yew woodland (Greater mouse-ear  Lowland beech & yew woodland  Improved grassland  Lowland meadow  Neutral grassland Myotis myotis)  Lowland wood-pasture & parkland  Machair  Wet woodland  Ancient &/or species rich hedgerows  Boundary & linear features  Aquifer fed naturally fluctuating water  Calcareous grassland bodies  Coniferous woodland  Chalk stream  Rivers & streams  Eutrophic standing waters  Standing open water & canals Barbastelle  Lowland beech & yew woodland  Lowland calcareous grassland Barbastella barbastellus  Lowland meadow  Lowland wood-pasture & parkland  Mesotrophic lakes  Native pine woodlands  Wet woodlands  Maritime cliffs & slopes Bechsteins  Ancient &/or species rich hedgerows  Boundary & linear features M. bechsteinii  Lowland beech & yew woodland  Broad-leaved & mixed yew  Lowland wood-pasture & parkland

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 Wet woodlands

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Appendix 2. A summary of habitat preferences compiled for various bat species studied using bat detector surveys in agricultural landscapes or widescale studies incorporating agricultural land classes.

Study area Study area character Species Habitats selected / Habitats avoided / Other habitats used Notes Source most used least used England, Wales & 32 British land classes All species combined Edge and linear Open and intensively Lowland unimproved Treelines selected in Scotland (910 1km in proportion to (Pipistrellus sp., habitats. Strong managed habitats. grassland only grass all land classes except squares) availability Myotis sp., Nyctalus preference for Openings in upland type not consistently two arable land types. (Walsh & Harris, noctula, Plecotus woodland edge and conifer plantations. avoided. 1996a) spp., Eptesicus water bodies. Broad- Arable land, improved NB Some habitats serotinus). leafed > mixed or grassland, semi- inconsistently selected conifer woodland. improved grassland. in different land class Woodland edge > types. woodland opening.

England, Scotland and 32 British land classes All species combined Woodland habitats Arable land. Relative abundance Wales in proportion to (Pipistrellus sp., (particularly lowest in intensively (Walsh & Harris, availability Myotis sp., Nyctalus broadleaf), treelines & farmed arable, 1996b) noctula, Plecotus hedgerows, riparian marginal upland and spp., Eptesicus (river, canal, lake and upland land classes serotinus). pond margin). towards the north of Britain. Significant negative gradient in abundance on south- north geographical axis.

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Appendix 2 continued. A summary of habitat preferences compiled for various bat species studied using bat detector surveys in agricultural landscapes or widescale studies incorporating agricultural land classes.

Study area Study area character Species Habitats selected / Habitats avoided / Other habitats used Notes Source most used least used Southwest England 30 study sites in Total bat activity 70% of all bat passes (Vaughan et al., undulating lowland recorded over rivers & 1997) pastoral landscapes lakes (sig. higher than covering 10 landuse in other landuse types types). Myotis and Plecotus Activity sig. greater High levels of activity group over rivers and lakes also recorded in than in other land use unimproved types. grassland, improved cattle pasture, ancient semi-natural woodland & mixed plantations. P. pipistrellus Rivers, lakes, Active and unimproved widespread across all grassland, amenity land use types, grassland, improved foraging distributed cattle pasture, arable evenly across seasons land, villages, ancient and land use types. semi-natural woodland, conifer & mixed plantations P. pygmaeus Rivers and lakes Significantly lower occurrence and foraging across other 8 land use types.

32 Project DEFRA title project code Appendix 2 continued. A summary of habitat preferences compiled for various bat species studied using bat detector surveys in agricultural landscapes or widescale studies incorporating agricultural land classes.

Study area Study area character Species Habitats selected / Habitats avoided / Other habitats used Notes Source most used least used Southwest England Nyctalus noctula Lakes, improved No sig. difference in Also recorded over (Vaughan et al., pasture. activity over ancient woodland and 1997) continued improved pasture, coniferous and mixed rivers, unimproved plantations. grassland, amenity grassland, arable land and villages. Nyctalus leisleri Lakes, rivers, Unimproved and improved pasture. amenity grassland, arable land, villages, ancient woodland coniferous and mixed plantations. Yorkshire Dales 13km upland stretch Myotis daubentonii River stretches with Rapid or cluttered National Park, of River Wharfe smooth water surface water with trees one England bordered by and trees on both side, smooth water (Warren et al., 2000) unimproved/semi- banks. with no trees. improved pasture and Pipistrellus Smooth water with Avoided rapid water Authors draw hay meadow. pipistrellus trees on both banks with no trees, rapid attention to potential Moorland lies above water with trees on importance of riparian valley. one side, smooth habitats in upland water with no trees systems where earlier evening. Later alternative foraging evening avoided rapid habitat may be or cluttered water infrequent. with trees on one side.

33 Project DEFRA title project code Appendix 2 continued. A summary of habitat preferences compiled for various bat species studied using bat detector surveys in agricultural landscapes or widescale studies incorporating agricultural land classes.

Study area Study area character Species Habitats selected / Habitats avoided / Other habitats used Notes Source most used least used Southwest England Paired river/stream Total bat activity Upstream of effluent Downstream of Downstream activity (Vaughan et al., stretches upstream source effluent source reduced by 11%. 1996) and downstream of 19 Downstream prey sewage treatment capture attempts works. reduced by 28%. Nyctalus spp. and No significant Eptesicus difference in serotinus group upstream/downstream activity. Myotis spp. No significant difference in upstream/downstream activity. No significant difference in feeding buzz counts. Downstream buzz ratios sig. increased by 112%. P. pipistrellus Upstream of effluent Downstream of Downstream activity source effluent source reduced by 55%. Feeding buzz counts reduced by 87% downstream. P. pygmaeus Upstream of effluent Downstream of Downstream activity source effluent source reduced by 51%. No significant difference in feeding buzz counts.

34 Project DEFRA title project code Appendix 2 continued. A summary of habitat preferences compiled for various bat species studied using bat detector surveys in agricultural landscapes or widescale studies incorporating agricultural land classes.

Study area Study area character Species Habitats selected / Habitats avoided / Other habitats used Notes Source most used least used Lower Derwent New woodland Overall bat activity New plantations Arable land Valley, East & North plantations and (comprising P. Yorkshire adjacent farmland in pipistrellus, P. (Moore et al., pastoral landscape. pygmaeus, Myotis (submitted)) spp., Nyctalus noctula) Vale of York 78 new woodland Pipistrellus species New plantations Arable land Significantly higher (Moore, 2002) plantations and combined activity over new adjacent farmland in plantations in mixed farming comparison of 2 landcape. habitats Nyctalus noctula No significant activity difference between 2 habitat types Myotis spp. No significant activity difference between 2 habitat types

35 Project DEFRA title project code Appendix 2 continued. A summary of habitat preferences compiled for various bat species studied using bat detector surveys in agricultural landscapes or widescale studies incorporating agricultural land classes.

Study area Study area character Species Habitats selected / Habitats avoided / Other habitats used Notes Source most used least used Northern Ireland (96 Eight major Northern Pipistrellus Treelines, Scattered/isolated Young/felled Selected and avoided 1km squares) Ireland land classes pipistrellus rivers/canals trees, hedgerows, plantations, deciduous fewer habitats than P. (Russ & Montgomery, represented. Northern open ditches, woodland centre & pygmaeus, found in 2002) Ireland approx. 63% streetlights, urban edge, mixed large number of agricultural land. buildings woodland centre & habitats. edge, coniferous woodland centre & edge, pond margins, lake/reservoir margins, covered ditches, improved grassland, lowland or upland-unimproved grassland, parkland/amenity grassland, arable land, streams, rural buildings Pipistrellus pygmaeus Deciduous woodland Improved grassland, Mixed woodland centre & edge, upland or lowland centre & edge, lake/reservoir margin, unimproved coniferous woodland river/canal grassland, centre & edge, pond scattered/isolated margin, trees, hedgerows, parkland/amenity open ditches, urban grassland, treelines, buildings streams, rural buildings

36 Project DEFRA title project code Appendix 2 continued. A summary of habitat preferences compiled for various bat species studied using bat detector surveys in agricultural landscapes or widescale studies incorporating agricultural land classes.

Study area Study area character Species Habitats selected / Habitats avoided / Other habitats used Notes Source most used least used Northern Ireland (96 Nyctalus leisleri Parkland/amenity Improved grassland, Young/felled Among studied 1km squares) grassland, deciduous hedgerow. plantation, deciduous species only one to (Russ & Montgomery, woodland edge, woodland centre, select open 2002) continued river/canal pond margin, parkland/amenity lake/reservoir margin, grassland. upland or lowland unimproved grassland, arable land, mixed woodland edge, coniferous woodland edge, treelines, scattered/isolated trees, streetlights, urban buildings Myotis group Deciduous woodland Improved grassland, Deciduous woodland centre, lake/reservoir upland or lowland centre, mixed margin, river/canal unimproved woodland centre & grassland, arable land, edge, coniferous coniferous woodland woodland centre, edge, pond margin, water scattered/isolated margins with no trees, hedgerows, vegetation, parkland/amenity grassland, treelines, streams, rural buildings

37 Project DEFRA title project code Appendix 2 continued. A summary of habitat preferences compiled for various bat species studied using bat detector surveys in agricultural landscapes or widescale studies incorporating agricultural land classes.

Study area Study area character Species Habitats selected / Habitats avoided / Other habitats used Notes Source most used least used Northern Ireland (96 All species including Deciduous woodland Marshland, heath, Young/felled Seven selected 1km squares) Pipistrellus nathusii centres & edge, mixed improved grassland, plantations, mixed habitats = water / (Russ & Montgomery, and Plecotus auritus woodland edge, upland-unimproved woodland centre, woodland types, 2002) continued treelines, pond grassland, coniferous woodland avoided = open areas margins, scattered/isolated centre & edge, or linear features not lake/reservoir trees, hedgerows, lowland-unimproved associated with large margins, river/canal open ditches, water grassland, amounts of margins with no parkland/amenity vegetation. Water vegetation grassland, arable land, bodies with no open stone wall, vegetation usually streams, rural avoided. buildings County Wexford, 5km radius of Nyctalus leisleri Limited evidence of Ireland Baldwinstown. habitat preference, (Shiel & Fairley, Lowland, improved except for streetlights. 1998) pastoral land. Well- developed hedges, no woodland. County Clare, Ireland 1km radius of farm Rhinolophus Greatest foraging Rarely observed over Trees, hedgerows. Use of farmyards may (McAney & Fairley, building roosts at hipposideros intensity at pasture. be related to decaying 1988) Ballynacally & waterbodies and organic matter, with Newhall. Permanent farmyards. potential local pasture, hedgerows, concentration of occasional woodland. Nematocera.

38 Project DEFRA title project code Appendix 3. A summary of foraging distances and foraging preferences compiled for species studied within an agricultural landscape by radiotracking, following capture and tagging at roost site. * denotes bats marked with reflective tape (not radio-tagged). # denotes findings based on single radio-tracked individual supported by systematic bat detector work.  denotes study based on single radio-tracked individual. MCP denotes minimum convex polygon.

Species Study area Study area character Home range / direct Preferred foraging Habitat least used Other habitat used distance travelled habitat Source Common pipistrelle Avon valley, Pastoral landscape, Total MCP for 12 bats Wide habitat range. Pipistrellus pipistrellus Hampshire, southern improved grazing = 4.4 km2. Maximum Trees in parkland, NB pregnant females England land, flood plain wet direct distance from trees in pasture, Davidson-Watts grassland and relict roost = 3km. One bat woodland blocks, (unpublished) watermeadows, followed for 17.8km mature trees occasional arable, in one night. overhanging arable. Soprano pipistrelle bordered by Total MCP for 24 Water bodies lined Pipistrellus pygmaeus deciduous forest. pregnant /lactating with trees. NB pregnant / lactating bats = 7.3 km2. females Maximum distance from roost = 2.3km. One bat followed for 8.5km in one night. Pipistrelle *Inverurie, Grampian Inverurie: lowland Inverurie: max. Inverurie: foraged Inverurie: No bats undifferentiated and *Drumore, agricultural river distance = 5.1km, over river or in thick seen foraging where Pipistrellus sp. Tayside, north-east valley comprising 3.7km (pre-, post- adjacent vegetation. river crossed Scotland urban, parkland, parturition). Average open fields with no NB pregnant / lactating (Racey & Swift, 1985) pasture, arable & distance = 1.8km pre- riparian trees. Not females conifer plantations. parturition, 1.3km observed in /near post parturition. conifer plantations.

Drumore: upland, Drumore: Max. Drumore: foraged in Drumore: Avoided loch with conifer distance before / post all parts with trees or open fields and hills. plantations, deciduous parturition =2.5km. water. trees, open hills. Average distance = 1.0km.

39 Project DEFRA title project code Appendix 3 continued. A summary of foraging distances and foraging preferences compiled for species studied within an agricultural landscape by radiotracking, following capture and tagging at roost site. * denotes bats marked with reflective tape (not radio-tagged). # denotes findings based on single radio-tracked individual supported by systematic bat detector work.  denotes study based on single radio-tracked individual. MCP denotes minimum convex polygon.

Species Study area Study area character Home range / direct Preferred foraging Habitat least used Other habitat used distance travelled habitat Source Daubenton’s Myotis * Spey valley, north- Agricultural river Pools, drainage daubentoni east Scotland valley ditches, river and (Swift & Racey, 1983) riparian vegetation. Whiskered Myotis Lower Derwent New woodland Maximum distance Woodlands, wetlands, Strong avoidance of mystacinus valley, East & North plantations and from roost adults = hedgerows, farmyards open areas. Yorkshire adjacent farmland in 3.5km (juvenile (Moore et al., pastoral landscape. closer). (submitted); Moore & Hutson, ??) Serotine Eptesicus Brighton & Lewes, Brighton: urban, Mean distance roost Brighton: white serotinus southern England parkland, amenity to feeding site = 4km, streetlamps, pasture, grassland, cattle- max. = 12km. downland, parkland. (Catto et al., 1996) grazed pasture, arable, Intensive feeding woodland & chalk recorded at large cow grassland. dung pile on farm. Lewes: lowland Lewes: chalkland pasture, chalk scrub and grassland, arable and watermeadow. woodland. ― ― ― ― ― ― south Mostly lowland arable Colony home range Pasture and Arable/ urban Cambridgeshire, and grazing farmland, 24 - 77km2. Colony woodland. Seasonal eastern England with river valleys and core area range 13 - variation (May, June, (Robinson & deciduous woodland. 33km2. Highly July > 53% foraging Stebbings, 1997) variable individual in woodland reducing MCP 0.16 – 47.6km2. to 40.5% and 28.1% in August and September.

40 Project DEFRA title project code Appendix 3 continued. A summary of foraging distances and foraging preferences compiled for species studied within an agricultural landscape by radiotracking, following capture and tagging at roost site. * denotes bats marked with reflective tape (not radio-tagged). # denotes findings based on single radio-tracked individual supported by systematic bat detector work.  denotes study based on single radio-tracked individual. MCP denotes minimum convex polygon.

Species Study area Study area character Home range / direct Preferred foraging Habitat least used Other habitat used distance travelled habitat Source Serotine Eptesicus south Maximum direct Simultaneous increase serotinus (continued) Cambridgeshire, distance to feeding in pasture foraging eastern England site =7.4km. from 49.8 to 63.4%). (Robinson & Stebbings, 1997) (continued) Leisler’s bat Nyctalus County Wexford, Mainly improved Maximum straight Two-thirds of Lights, estuary, leisleri Ireland agricultural land line distance from recorded foraging stream, beach and dominated by pasture, roost in 1 night = time over pasture or dunes, forest. (Shiel et al., 1999) plus localised area of 13.4km. Mean falls drainage canals. mountain conifer from preparturition to forest. lactation. ― ― ― ― ― ― Kent: rural matrix of Mean maximum Kent: woodland & Kent: urban, then Bristol, south-west woodland, urban, distance from roost = roads sig. > arable sig. arable. England & north arable, ungrazed 4.2km, range 2.88 - > urban. Kent, south-east grassland & roads; 5.75km. Average Bristol: greater Bristol: urban and woodland, amenity England Bristol: city centre, individual MCP = variability in arable. grassland, riparian (Waters et al., 1999) urban, pasture, arable, 7.4km2, range MCP = preferences but amenity grass, 2.42-18.36km2. pasture significantly parkland, riparian, preferred to amenity woodland. grassland, arable or urban.

41 Project DEFRA title project code Appendix 3 continued. A summary of foraging distances and foraging preferences compiled for species studied within an agricultural landscape by radiotracking, following capture and tagging at roost site. * denotes bats marked with reflective tape (not radio-tagged). # denotes findings based on single radio-tracked individual supported by systematic bat detector work.  denotes study based on single radio-tracked individual. MCP denotes minimum convex polygon.

Species Study area Study area character Home range / direct Preferred foraging Habitat least used Other habitat used distance travelled habitat Source Brown long-eared north-east Scotland Agricultural land Maximum distance 42% of foraging sites 9% foraging sites Native non-plantation Plecotus auritus (Entwistle et al., (pasture and arable), from roost =2.8km =mature deciduous close to water. conifers, birch 1996) woodland, villages, (male). 92% of time woodland. Mixed woodland. Edges of water bodies. bats within 1.5km of woodland. non-native conifer roost. Females spent stands. 17% foraging sig. more time < sites trees adjoining 0.5km and males sig. pasture. more time >1.5km from roost. ― ― ― ― ― ― * Spey valley, north- Agricultural river Maximum distance Coniferous and Isolated trees east Scotland valley from roost = 1.1km deciduous woodland (Swift & Racey, 1983) ― ― ― ― ― ― Lower Derwent New woodland Maximum distance Broad-leaved Arable land, open Coniferous woodland valley, East & North plantations and from roost adults = woodland including land Yorkshire adjacent farmland in 7.4km, juveniles = single mature trees, (Moore et al., pastoral landscape. 2.7km hedge trees, parkland (submitted)) trees & small woodland Lesser horseshoe Monmouthshire, Traditional pastoral Foraging ranges 12- Woodland Pasture and arable Rhinolophus south Wales farming landscape 53 ha (100% kernel). (predominantly least used by foraging hipposideros (Bontadina et al., predominant, plus 50% foraging activity broadleaf). bats. 2002) some intensively within 600m of NB habitat diversity > managed agricultural maternity roost. in foraging areas used land. Maximum distance than in available from roost 4.2km. colony range.

42 Project DEFRA title project code Appendix 3 continued. A summary of foraging distances and foraging preferences compiled for species studied within an agricultural landscape by radiotracking, following capture and tagging at roost site. * denotes bats marked with reflective tape (not radio-tagged). # denotes findings based on single radio-tracked individual supported by systematic bat detector work.  denotes study based on single radio-tracked individual. MCP denotes minimum convex polygon.

Species Study area Study area character Home range / direct Preferred foraging Habitat least used Other habitat used distance travelled habitat Source Lesser horseshoe Ciliau, Radnor, Wales Pastoral farm Individual MCP range Riparian broadleaf Individuals recorded Rhinolophus (Schofield et al., landscape with 11.84-353.00 ha woodland. foraging in conifer hipposideros (continued) 2002) lowland cattle grazing (mean 96.74 ha). plantation. and upland sheep Total colony MCP grazing. Broadleaf 851.70ha. Core woodland, including foraging area 8.33- riparian woodland, 34.68ha (mean conifer plantations. 20.67ha). Maximum distance travelled range 0.440-2.695 (mean 1.4km). ― ― ― ― ― ― # Revogne, Famenne, Village surrounded by Maximum observed Deciduous woodland Arable and spruce Some foraging in tall Belgium permanent grazing distance from roost with copses or mixed woodland avoided. hedgerows connecting (Motte & Libois, meadows, dense 1.2km (assumed less coniferous woodland. woodland foraging 2002) hedgerow network, than actual). areas. deciduous or mixed pine woods on nearby hilltops. ― ― ― ― ― ―  Herreninsel, Agricultural land , Max recorded Main foraging areas Hedges, tree lines Park/garden with Bavaria, Germany roads, gardens, foraging distance were mixed forests. streams, never used trees. (Holzhaider et al., houses, orchards, from roost 3.6km for foraging. 2002) coniferous and (mean 2.4km). deciduous or mixed forests

43 Project DEFRA title project code

Appendix 3 continued. A summary of foraging distances and foraging preferences compiled for species studied within an agricultural landscape by radiotracking, following capture and tagging at roost site. * denotes bats marked with reflective tape (not radio-tagged). # denotes findings based on single radio-tracked individual supported by systematic bat detector work.  denotes study based on single radio-tracked individual. MCP denotes minimum convex polygon.

Species Study area Study area character Home range / direct Preferred foraging Habitat least used Other habitat used distance travelled habitat Source Greater horseshoe King’s Wood and c. 14 land types Adults Spring: ancient Rhinolophus Woodchester Park, including woodland, semi-natural Adults ferrumequinum south-west England parkland, urban, woodland & cattle- Spring: Least (Duvergé & Jones, scrub, amenity, grazed pasture, then important habitats = 2003) arable, pasture and sheep/horse-grazed coniferous woodland, hay/silage grassland. pasture & meadow. arable and urban. Summer & autumn: cattle-grazed pasture Summer & autumn: then ancient Least important = woodland, cattle- arable and urban grazed meadow & sheep/horse-grazed pasture. Juveniles: Cattle-grazed pastures Juveniles Arable and urban

44 Project DEFRA title project code Appendix 4. Summary of the ecological traits major insects families occurring in the diet of bats (Vaughan, 1997; Rydell & Petersons, 1998; Arlettaz et al., 2000; Rostovskaya et al., 2000) and the potential impacts of farming. Key to adult activity: N = nocturnal, C = crepuscular and D = diurnal. Invertebrate ecology sources (Colyer & Hammond, 1951; Smith, 1989; Emmet & Heath, 1991; Chinery, 1993)

Order Family Common Adult Habitat/behaviour of Principle habitats of larvae Potential impacts of farming Sub-order name activity adults Diptera Tipulidae Crane-fly, N (mostly) Grassland, swarming Damp woodlands, decaying wood, 1. Pesticides leatherjacket common in smaller species leaf litter, grasslands/cereal (pests 2. Drainage/irrigation Nematocera feed on roots), some ponds & 3. Ditch management streams, 4. Woodland management 5. Tillage 6. FYM/slurry/Compost Diptera Chironomidae Non-biting C Males swarm over water Mainly aquatic often dominant, 1. Pesticide run off. midges bodies, over trees or bushes some decaying matter (terrestrial) in 2. Drainage/irrigation Nematocera open vegetation (i.e. early 3. Dredging. succession) 4. FYM/slurry/Compost 5. Set aside 6. Tillage Diptera Anisophodidae Window C Some species swarm, damp Rotting vegetables, damp woodland 1. Woodland management midges woodland, near damp 2. Drainage/irrigation Nematocera decaying matter (vegetables) Diptera Culicidae Mosquitoes N Near water Fresh, fouled or brackish water, salt 1. Pesticide run off. marshes, tree hole and most other 2. Drainage/irrigation Nematocera inland ground water, including 3. Dredging sewage Diptera Mycetopilidae Fungus gnats C Damp woodland, fungi Fungi, usually on rotting wood, 1. Woodland management some grass tussocks or ground Nematocera fungi Diptera Ceratopogonidae Biting midges C Near water (as larvae), Mainly fresh water, streams, lakes, 1. Pesticide run off. around buildings ponds, swamps & ditches usually 2. Drainage/irrigation Nematocera with plenty of organic matter, some 3. Dredging terrestrial or semi-aquatic, e.g. in liquid products of farmyard heaps Diptera Simulidae Black-flies C? Near water Running water 1. Pesticide run off. 2. Drainage/irrigation Nematocera 3. Dredging

45 Project DEFRA title project code Appendix 4 continued. Summary of the ecological traits major insects families occurring in the diet of bats (Vaughan, 1997; Rydell & Petersons, 1998; Arlettaz et al., 2000; Rostovskaya et al., 2000) and the potential impacts of farming. Key to adult activity: N = nocturnal, C = crepuscular and D = diurnal. Invertebrate ecology sources (Colyer & Hammond, 1951; Smith, 1989; Emmet & Heath, 1991; Chinery, 1993)

Order Family Common Adult Habitat/behaviour of Principle habitats of larvae Potential impacts of farming Sub-order name activity adults Diptera Sciaridae D? Often in buildings Rotting matter, fungi (pests of 1. Pesticides mushrooms) 2. FYM/slurry/Compost Nematocera Diptera Psychodidae Owl midges C Form huge swarms at Decaying matter, usually in water, 1. Pesticide run off breeding sites e.g. sewage filter beds. 2. Drainage Nematocera Diptera Bibionidae St Marks fly D Woodland & grassland, can Soil (woodland & open country), 1. Pesticides etc. be very dense in spring dung & wood debris , (possible pest 2. Drainage/irrigation Nematocera as eat roots) 3. Fertiliser Diptera Cecidomyiidae Gall midges Swarm over lakes and ponds Decaying plant material, in soil, 1. Pesticides some gall forming (Hessian fly pest 2. Tillage Nematocera of wheat though not much in UK, also Pear midge) others predatory Diptera Trichoceridae Winter gnats D, C Abundant in spring/autumn, Leaf mould, fungi and decaying 1. Pesticides cool shady places in summer vegetable matter 2. Farm husbandry Nematocera Diptera Muscidae Incl. House D Around dung and animal Decaying material 1. Pesticides/avermectins fly, Stable fly housing and rubbish tips 2. Farm husbandry Diptera Empidae Dance flies D Congregate in swarms, often Soil, fungi etc. in woods, wet 1. Pesticides over water, visit flowers meadows, mosses & algae in rivers, 2. Loss of flower rich areas streams & lakes 3. Dredging?? Diptera Calliphoridae Incl. D Animal corpses, around Animal corpses 1. Farm husbandry Blue/green buildings bottles Diptera Syrphidae Hover flies D Many require flower rich Wide range of habitats, some 1. Pesticides. areas (pollen feeders) associated with aphids, rotting 2. Loss of flower rich areas wood, leaf miners, compost heaps, some aquatic especially stagnant water especially around farmyards Diptera Stratiomyiidae (Incl. soldier D On waterside vegetation, Some aquatic, leaf litter rotting 1. Loss of hedgerows. flies) especially coastal organic matter, dung, some found in 2. Pesticide spray drift. hedgerows in moss/leaf litter 3. Use of compost/FYM

46 Project DEFRA title project code Appendix 4 continued. Summary of the ecological traits major insects families occurring in the diet of bats (Vaughan, 1997; Rydell & Petersons, 1998; Arlettaz et al., 2000; Rostovskaya et al., 2000) and the potential impacts of farming. Key to adult activity: N = nocturnal, C = crepuscular and D = diurnal. Invertebrate ecology sources (Colyer & Hammond, 1951; Smith, 1989; Emmet & Heath, 1991; Chinery, 1993)

Order Family Common Adult Habitat/behaviour of Principle habitats of larvae Potential impacts of farming Sub-order name activity adults Diptera Sphaeroceridae D Around dung? Dung 1. Livestock management. 2. Avermectins Diptera Rhagionidae Snipe flies Some damp shady places, Soil, leaf litter, rotting wood 1. Pesticides bracken, streamside 2. Tillage vegetation etc. Diptera Dolichopodidae D Damp places near water, Damp soil, sand, rotting wood, 1. Water management some can be very abundant under bark, sap, cattle dung, plant over ponds etc. stems & some aquatic. Diptera Ephydridae Shore flies Sea shore and edges of Sea shore and edges of lakes/ponds, 1. Pesticide spray drift. lakes/ponds leaf miners, predators, detritivores 2. Management of riparian vegetation. 3. Dredging. Diptera Asilidae Robber flies D Often grasslands Dry habitats, sandy soils in 1. Rolling/harrowing pastures grasslands, woods, dunes, some 2. Pesticides dead vegetable matter & dung Diptera Fanniidae Incl. Lesser D? Around dung and buildings Dung 1. Livestock management. housefly 2. Avermectins Diptera Phoridae Scuttle flies D On vegetation, enter Very wide range of habitats 1. Pesticides buildings including decaying material, dead 2. Farm husbandry animals, (some pests of mushrooms) Diptera Tachinidae D Often around flowers and Parasitic mainly on butterflies and 1. Pesticides near hosts moths Diptera Lauxiniidae Shady vegetation in damp Decaying vegetation especially 1. Pesticides places leaves & straw, some attack plants 2. Drainage 3. Hedgerow removal? Diptera Opomyzidae Grasslands, cereals, Shoots of grasses incl. cereals 1. Pesticides. woodlands Diptera Sarcophagidae Flesh flies D Near animal corpses Animal corpses 1. Farm husbandry Diptera Clusiidae D Woodland Rotting wood 1. Woodland management

47 Project DEFRA title project code Appendix 4 continued. Summary of the ecological traits major insects families occurring in the diet of bats (Vaughan, 1997; Rydell & Petersons, 1998; Arlettaz et al., 2000; Rostovskaya et al., 2000) and the potential impacts of farming. Key to adult activity: N = nocturnal, C = crepuscular and D = diurnal. Invertebrate ecology sources (Colyer & Hammond, 1951; Smith, 1989; Emmet & Heath, 1991; Chinery, 1993)

Order Family Common Adult Habitat/behaviour of Principle habitats of larvae Potential impacts of farming Sub-order name activity adults Diptera Tabanidae Horse flies, D Around livestock Damp soil or sand adjoining water 1. Pesticides gad flies, bodies, some aquatic and some dry 2. Drainage/irrigation stouts, clegs meadows. Diptera Chloropidae Frit fly D? Grasses & low vegetation in Vegetation, some crop pests 1. Pesticides meadows, roadside verges, 2. Cutting/grazing waste land Diptera Helomyzidae? D Around decomposing animal Decomposing animal or vegetable 1. Farm husbandry? or vegetable matter matter in woods, rabbit burrows, caves with bats, cess-pits Lepidoptera Noctuidae Mainly N Wide range Can be crop pests, e.g. cutworms 1. Pesticides Lepidoptera Geometridae Mainly N Wide range Soil or herbage 1. Pesticides 2. Hedgerow management 3. Tillage Lepidoptera Sphingidae Hawkmoths N/C some Where foodplants occur, Feed on foliage of trees 1. Woodland management diurnal often in woodlands Lepidoptera Arctiidae e.g. cinnibar, Mainly N Wide range of habitats, Low plants, especially lichens 1. Pesticides tiger mostly near lichen 2. Woodland management Lepidoptera Hepialidae C Mostly grasslands, some in Roots of grasses, dandelions, 1. Pesticides woodland bracken 2. Rolling/harrowing Lepidoptera Lasiocampidae Mainly N, Woodlands, grasslands, Some (lackey's) build tents in 1. Pesticides some males heathland shrubs & trees, pest in apple 2. Hedge management D orchards Lepidoptera Lymantriidae Tussock moths Mainly N Woodland, grassland, Mostly on tree foliage, some pests 1. Pesticides wetland, coastal on fruit trees 2. Woodland management Lepidoptera Notodontidae N Wide range Exclusively on tees foliage 1. Pesticides 2. Woodland management Lepidoptera Nymphalidae e.g. D Around host plant, Wide range of plants including 1. Pesticides Tortoiseshell, hedgerows etc. nettles, grasses etc. 2. Weed control fritillaries Lepidoptera Pyralidae N/C/D Wide range of habitats Mostly feed on grasses, some grain 1. Pesticides & other stored products, pests on 2. Cutting/grazing cereals.

48 Project DEFRA title project code

Lepidoptera Drepanidae Mainly N Mainly woodland, On tree foliage 1. Woodland management

49 Project DEFRA title project code Appendix 4 continued. Summary of the ecological traits major insects families occurring in the diet of bats (Vaughan, 1997; Rydell & Petersons, 1998; Arlettaz et al., 2000; Rostovskaya et al., 2000) and the potential impacts of farming. Key to adult activity: N = nocturnal, C = crepuscular and D = diurnal. Invertebrate ecology sources (Colyer & Hammond, 1951; Smith, 1989; Emmet & Heath, 1991; Chinery, 1993)

Order Family Common Adult Habitat/behaviour of Principle habitats of larvae Potential impacts of farming Sub-order name activity adults Lepidoptera Endromidae Males D/C Woodland Birch, alder only Scotland 1. Woodland management Females N Lepidoptera Yponomuetidae N/C/D Hedges and small trees Hedges and small trees 1. Hedge management 2. Woodland management Lepidoptera Zygaenidae Burnets & D Wide range Wide range 1. Pesticides foresters Lepidoptera Thyatiridae Mainly N, Mainly woodlands Trees and brambles, hide in spun 1. Pesticides some also C leaves by day, emerge at night 2. Woodland management Coleoptera Carabidae Ground N and D Agricultural fields, Many in soil and grassy tussocks 3. Pesticides beetles grassland, woodland, hedges 4. Tillage etc. Smaller species fly some 5. Cutting/grazing at night, other larger specie 6. Hedgerow management surface active at night or day. Coleoptera Scarabaeidae Dung beetles, D and N Some species e.g. May bug Dung, some dead wood and plants 1. Avermectins chafers fly at night 2. Pesticides 3. Farm husbandry Coleoptera Geotrupidae Dor beetles D and N Large species fly at night Dung 1. Avermectins 2. Pesticides 3. Farm husbandry Coleoptera Silphidae Burying N? Attracted to animal corpses Carrion and some in fungi 1. Farm husbandry beetles, Sexton beetles Coleoptera Cerambycidae Long-horn D and N Some species fly at night, Dead wood 1. Woodland management beetles need flowers, (umbellifers, 2. Loss of flower rich areas?? hawthorn etc.) Coleoptera Curculionidae Weevils D On vegetation, especially Plant roots, stems, leaves, several 1. Pesticides forbs. pest species on crops 2. Cutting/grazing

50 Project DEFRA title project code Appendix 4 continued. Summary of the ecological traits major insects families occurring in the diet of bats (Vaughan, 1997; Rydell & Petersons, 1998; Arlettaz et al., 2000; Rostovskaya et al., 2000) and the potential impacts of farming. Key to adult activity: N = nocturnal, C = crepuscular and D = diurnal. Invertebrate ecology sources (Colyer & Hammond, 1951; Smith, 1989; Emmet & Heath, 1991; Chinery, 1993)

Order Family Common Adult Habitat/behaviour of Principle habitats of larvae Potential impacts of farming Sub-order name activity adults Coleoptera Elateridae Click beetles D and N Some species fly at night Dead wood, plants and in soil, some 1. Pesticides pests of crops (e.g. wireworms) Ephemeroptera Ephemeridae Mayflies C, N Near water, weak fliers, Aquatic, slow moving or still 1. Pesticide run off many emerge in the evening water, with muddy bottom 2. Dredging Ephemeroptera Caenidae and die by morning, males Bottom dwellers Ephemeroptera Baetidae swarm over water Swimmers, some running water other ponds & canals Neuroptera Hemerobiidae Brown Mostly N?? Mostly woodlands Aphid predators, mostly woodland 1. Pesticides lacewings (especially pine trees) 2. Woodland management

Hemiptera Unknown True bugs, Mainly D, N Mainly plant feeding in wide As adults 1. Pesticides aphids, range of habitats 2. Cutting/grazing leafhoppers 3. Hedgerow management Tricoptera Unknown Caddis flies C Normally near water, large Aquatic 1. Pesticide runoff numbers can occur during 2. Riparian management emergence 3. Dredging Arachnida Unknown Spiders and N & D Active hunters on ground, Varied 1. Pesticides allies vegetation, passive hunters - 2. Cutting/grazing orb weavers need structured vegetation Hymenoptera Unknown Wasps, bees, D Many reliant on flower rich Warm sunny areas with bare earth, 1. Pesticides sawflies, ants (occasionally habitats for nectar & pollen. vegetation for plant feeders such as 2. Loss of lower rich areas etc N) Some species of wasps fly at sawflies. night Psocoptera Unknown Book lice ?? Herbage and trees As adults, often under bark & in 1. Pesticides bird nests 2. Woodland management Dermaptera Unknown Earwigs N Ubiquitous As adults, early stages in soil 1. Pesticides 2. Tillage Plecoptera Unknown Stoneflies D? Streamside vegetation & Aquatic, mostly hill streams and 1. Pesticide run off stones, only fly weakly chalk streams 2. Dredging 3. Riparian management

51 Project DEFRA title project code Appendix 5. Summary of agri-environment scheme prescriptions of significance to bats in agricultural land. Underlined text = reference to mammals/insect eating animals. Bold text = reference to bats. Key to scheme acronyms: ESA = Environmentally Sensitive Areas, CSS = Countryside Stewardship Scheme, CSM = Countryside Management Scheme and RSS = Rural Stewardship Scheme.

Country Scheme Hedge Grassland Riparian Trees Pesticide regulation  No removal  Arable reversion  Pond creation &  Retain & maintain  Weed control  Maintain all  Hay meadows restoration woodland  Permanent grassland  Restoration  Wet meadows  Buffer strips  Pollarding  Arable margin buffer  Planting  Limited fertilisers  Maintain ditches &  Planting strip dykes  Restoring traditional  Unfertilised ESA  Laying  Extensification  Coppicing  Rough grassland  Water levels orchards headlands  Hedge bank restoration  Wilt & turn silage  Streamside margins  Hedgerow trees  Arable reversion  Buffer strips  Buffer zones  Uncropped wildlife strips  Clean land England  Laying  Buffer strips  Bankside  Planting (single or  Conservation  Planting  Wildlife strips management blocks) headlands  Coppicing  Park restoration  Buffer strips  Coppicing  Buffer zones  Restoring banks  Hay meadows  Hay meadows  Pollarding  Recreating meadows  Grazed pastures  Old orchards CSS  Grazed meadows  Water meadow  Arable 6m margins restoration  Beetles banks & 2m  Water level control margins  Reedbeds  Fen  Ponds & scrapes  Retain hedges  Unimproved  Buffer strip  Retain individual  1 m buffer strips  25 % uncut each year grassland  Establish streamside trees  Unsprayed crops  Repair/establish  Rough grass margin corridors  Broad-leaved Wales Tir Gofal boundaries  Convert arable to  Creating ponds woodland grassland  establishment  Grassland restoration  Traditional orchards  Parkland

52 Project DEFRA title project code Appendix 5 continued. Summary of agri-environment scheme prescriptions of significance to bats in agricultural land. Underlined text = reference to mammals/insect eating animals. Bold text = reference to bats. Key to scheme acronyms: ESA = Environmentally Sensitive Areas, CSS = Countryside Stewardship Scheme, CSM = Countryside Management Scheme and RSS = Rural Stewardship Scheme.

Country Scheme Hedge Grassland Riparian Trees Pesticide regulation  Extended hedges  Species rich  Water margin  Plant along water  Grass margins  Hedge management grasslands management margins  Conservation  Planting  Management  Ponds  Native/semi-natural headlands  Laying  Creation  Creation woodland  Hedge management Scotland RSS  coppicing  Wetland  Restoration  Management  Exclude stock  Creation  Plant trees  Grass margin & beetle banks  Field boundaries  Species rich  Wetlands  Farm woodland  Species rich grassland  Hedge restoration grassland  Land adjacent to  Parkland  Wetlands  Interplanting  Grazed lakes/rivers  Traditional orchards  Farm woodland  Laying  Hay meadow  Buffer strips  Buffer strips ESA  Coppicing  Rough grass margin  Adjacent fields  Parkland N. Ireland  Buffer strips  Rough grass margin  Conservation crop margin  Traditional orchards CMS As ESA As ESA As ESA As ESA As ESA

53 Project DEFRA title project code Appendix 6. Summary of prescriptions of significance to bats in agricultural land available within Environmentally Sensitive Areas in England. Underlined text = reference to mammals/insect eating animals. Bold text = reference to bats. Key to scheme acronyms:

Scheme Hedge Grassland Riparian Trees Pesticide regulation  No removal  Extensive grassland  Arable margin buffer  No removal of willows  Weed control Upper  Hedgerow restoration  Wet grassland strip  Retain broad-leaved  Permanent grassland Thames  Laying  Arable reversion  Pond creation woodland  Arable margin buffer Tributaries  Planting  Hay making  Willow pollarding strip  Coppicing  Unfertilised headlands  No removal  Limited fertiliser use  Dykes  Planting  Weed control  Maintain hedges  Arable reversion  No removal of ponds etc.  Pollarding  Arable grassland  Laying  Arable grassland  Wet grassland - no margins Broads  Planting margins fertilisers  Permanent grassland  Coppicing  Maintain water levels  Fen  Fens  Arable reversion  Pond creation  Maintain hedges  Limited fertiliser use  Woodland management  Meadow & pastures Pennine  Planting  Meadows plan Dales  Laying  Planting  Coppicing  No fertiliser within 5m  Arable reversion  Wet grassland  Pollarding willows  Permanent grassland  Laying  Arable margin buffer  Tree planting  Arable margin buffer Test & zone zone Avon  Planting  Pond creation & Valleys  Coppicing restoration  Protect ditches  Maintain hedges  No inorganic fertiliser  Pond creation &  Maintain woodland  Permanent grassland North Kent  Planting restoration  Buffer strips Marshes  Laying  Protect ditches  Coppicing  Buffer strips  All retained  Arable reversion  Maintain ditches & dyke  Maintain trees  Permanent grassland Suffolk  Buffer strips (incl. margins)  Pollarding  Buffer strips River  Planting  Buffer strips  Retain woodland Valleys  Laying  Pond creation &  Planting  Coppicing restoration

54 Project DEFRA title project code Appendix 6 continued. Summary of prescriptions of significance to bats in agricultural land available within Environmentally Sensitive Areas in England. Underlined text = reference to mammals/insect eating animals. Bold text = reference to bats. Key to scheme acronyms:

Scheme Hedge Grassland Riparian Trees Pesticide regulation  No removal  Meadows  Pond creation &  Maintain woodlands  Inbye  Maintain all restoration  Planting  Meadows  Restoration  Pollarding Lake  Planting  Restoration of traditional District  Laying orchards  Coppicing  Hedge bank restoration  No removal  Regeneration of  Pond creation &  Maintain woodlands  Permanent grassland  Maintain all extensive pastures restoration South West  Planting  Hay meadows Peak  Laying  Hay meadow  Coppicing regeneration  No removal  Limit fertiliser  Maintain margins of  Retain woodland  Low input permanent  Maintain all streams  Management plan grassland & unimproved  Restoration  Pond creation &  Restoration of traditional Exmoor  Bank restoration restoration orchards  Planting  Laying

 No removal  Hay meadows  Retain ponds & wet  Retain woodland  All land  Maintain all  Limited fertiliser areas North Peak  Restoration  Maintain streamsides  Planting  Pond creation &  Laying restoration  Restoration  Limited fertilisers  Rotation maintenance of  Hedgerow trees  Permanent grassland  Maintain  Hay meadows ditches  Maintain woodland  Conservation headlands  No removal  Reversion to extensive  Maintain ponds  Pollarding Clun  Replanting grassland  Pond creation &  Laying  Arable reversion restoration  Coppicing  River bank restoration

55 Project DEFRA title project code Appendix 6 continued. Summary of prescriptions of significance to bats in agricultural land available within Environmentally Sensitive Areas in England. Underlined text = reference to mammals/insect eating animals. Bold text = reference to bats. Key to scheme acronyms:

Scheme Hedge Grassland Riparian Trees Pesticide regulation  Maintain hedges  Arable reversion  Maintain ponds &  Maintain woodland  Uncropped wildlife strip  Coppicing pingos  Restore pine belts  Conservation headland  Pond creation & Breckland  River valley grasslands restoration  Dyke & ditch creation & restoration  No removal  Rough grassland  Maintain ponds &  Planting  Field margin & clean West  Maintain  No fertiliser streams land Penwith  Field margin (1m)  Pond creation  Rough grassland  Cornish hedges  Maintain  Hay meadows  Retain water features  Maintain woodland  Arable land (1m from  No removal  Limited fertiliser  Fertiliser buffer zones  Planting boundary)  No damage  Pond creation &  Traditional orchards  Low input permanent Dartmoor  Planting restoration grassland  Laying  Unimproved & rough  Coppicing grassland  Hay meadows  Maintain  Limited fertiliser  Maintain all water  Hedgerow trees  Grassland margins  Restore  Grassland field margin features including banks  Planting  All land (1m from  Planting  Hay meadows & margins  Traditional orchards boundary Blackdown  Grassland margins hills  Laying  Low input, unimproved  Coppicing  Fertiliser buffer zones & rough grassland  Hedgebank restoration  Pond creation &  Hay meadows restoration  Maintain  Limited fertiliser  Maintain water levels in  Maintain  Permanent grassland  Planting  Arable reversion ditches  Laying  Fertiliser buffer zones  Coppicing  Maintain ditches & dikes Essex Coast (incl. Margins)  Restore & create dykes & ditches  Pond creation & restoration

56 Project DEFRA title project code Appendix 6 continued. Summary of prescriptions of significance to bats in agricultural land available within Environmentally Sensitive Areas in England. Underlined text = reference to mammals/insect eating animals. Bold text = reference to bats. Key to scheme acronyms:

Scheme Hedge Grassland Riparian Trees Pesticide regulation  No removal  Arable reversion  Retain & maintain all  Maintain  Conservation headlands  Maintain  Limited fertiliser watercourses & their  Retain broad-leaved  Improved & extensive  Management plan margins & banks woodland permanent grassland Cotswold  Pond creation &  4m buffer zone on Hills  Planting  Laying restoration individual trees  Coppicing  Planting  Pollarding  No removal  Limited fertiliser  Fertiliser buffer zones  Maintain  Boundary buffer zone  Maintain  Hay meadow  Maintain watercourses &  Planting (1m) Shropshire  Planting  Wilt & turn silage before margins & banks  Provision & protection  Permanent grassland Hills  Coppicing removal  Pond creation & of hedgerow trees  Laying restoration

 Maintain  Wilt & turn silage  Maintain ditches &  Maintain  Field boundary buffer  Plant  Downland turf creation streams  Willow pollarding (2m)  Laying  Arable reversion  Retain ponds &  Low input & downland South dewponds grassland Wessex  Coppicing  Restore ditches  Arable reversion Downs  Pond creation &  Downland turf creation restoration  Conservation headlands  Restore dewponds  Planting  Chalk grassland  Maintain ditches  Planting  Chalk grassland  Laying  Arable reversion  Retain ponds &  Conservation headland  Coppicing  Chalk dewponds  Permanent grassland South Downs  Permanent  Pond creation & restoration  Restore & create ditches & dykes

57 Project DEFRA title project code

Appendix 7 Current practices in the delivery of agri-environment information to land managers and the implementation of conservation measures on farmland.

1 Introduction

A series of discussions with individuals involved in promoting conservation on farmland considered the general process by which land-managers are made aware of environmental issues and encouraged to incorporate measures that support biodiversity. Individuals involved at different stages of this process, from first contact with land-managers through delivery of environmental information, assessment of the environmental status of farms, application to agri-environment schemes, evaluation of applications, development of agreements and subsequent administration, identified factors that influence the degree to which individual species are considered.

Personnel associated with the delivery of agri-environment information are drawn from a wide range of backgrounds including agriculture, ecology, rural resource management, land agency, biological research, soil science, drainage engineering, and former employment with statutory nature conservation organisations. It was notable that in the majority of cases, the regions covered by FWAG advisors, SAC advisors and RDS project officers / biodiversity co- ordinators or ecologists were sufficiently large to incorporate broad variation in agricultural land type e.g. mixed farms and dairy/beef or arable specialists, lowland and upland, horticulture, conventional and organic, and both extensively and intensively managed systems. Individuals working in this sector are therefore usually required to have a broad knowledge base and the flexibility to tailor advice to the circumstances of individual land managers.

Although this review drew on the experiences of the FWAG, SAC, RDS and CCW network it is recognised that land managers also obtain environmental information from many other sources including land agents, ADAS, wildlife trust consultants, RSPB, independent ecologists and agronomists. Several of these organisations also act as partners to RDS in the delivery of the major agri-environment schemes (Countryside Stewardship, ESA etc see section 4), or provide general advice or facilitate entry to local initiatives.

2 Initial contact with land-managers

FWAG and SAC advisors reported similar experiences of first contact with land managers, and approaches to initial farm assessment were comparable. In the main, advisor response was reactive i.e. a response to an enquiry by a farmer. Many farmers are eligible for a free initial environmental consultation, funded by DEFRA or a statutory nature conservation organisation. The enquiry is often motivated by an interest in agri-environment schemes, but non-scheme related enquiries, such as queries relating to a specific feature or species are also received. Typically an initial consultation combines discussion with the land manager, examining the nature of the business and the land manager’s aspirations. A walk/drive of the farm is led by the manager’s interests, although most advisors encouraged a whole farm approach, and is an opportunity to do a basic appraisal of landscape and habitat character. After the visit, the land manager receives a short, personalised report outlining the existing and potential interests of the site, with suggested follow-up options. A high proportion of land managers were reported to contact the advisory organisation for further consultation, e.g. to gain assistance with agri-environment scheme application.

Just over half the advisors reported that bats were frequently discussed with land managers on farm visits, with the topic mainly initiated by the advisor. No geographical pattern in tendency to discuss bats was apparent and several individuals based in regions where the two horseshoe species occur were among those people who reported rare or infrequent discussions about bats. Discussions on bats dealt with specific information (e.g. farmer’s observations of bats, management for greater horseshoes etc.) but often used bats to illustrate the value of habitat management, such as maintaining the connectivity of linear features. Several advisors commented that if bats were not mentioned on a farm visit, they would often be included in the summary report in this way.

3 Implementing conservation measures

Advisors facilitating an application to an agri-environment scheme return to the farm to conduct an audit of key habitats. Limited time restricts the amount of information that can be collated on species to that observed by the advisor during the visit, known by the land manager or available through other sources such as distribution maps, biological recording centres, reported by SNCO’s etc. Scheme applications are assigned to an RDS project officer (in England) who scores the application on nationally agreed criteria, evaluating the existing and potential value of the four Countryside Stewardship scheme objectives (historical features, landscape, public access and wildlife), taking into

58 Project DEFRA title project code account the location of the land in a target area, proximity to designated sites, protected habitats or species. Similar application and scoring processes operate in Scotland, Wales and Northern Ireland. Applications above a threshold score are visited by the project officer and an agreement negotiated between the project officer, land manager and partner organisation, based on measures drawn from the range of options listed in the scheme (for examples of habitat measures of particular relevance to bats see Annexes 5 and 6). Where land managers make an independent application, the project officer’s visit may be the first contact between the land manager and an agri-environment professional.

In contrast to the network of FWAG and SAC advisors, the majority of RDS biodiversity co-ordinators /ecologists interviewed believed that discussions relating to bats occurred infrequently. The individual that reported frequent discussions with land managers was a bat specialist, located in a region where land management for greater horseshoe bats was promoted actively. The apparent disparity between the view of the two groups (FWAG/SAC advisors and RDS biodiversity co-ordinators/ecologists) may indicate a situation where bats are frequently discussed on farm visits, or listed in reports, as species likely to benefit from habitat management, but seldom drive decisions to instigate particular land management. RDS biodiversity co-ordinators/ecologists, who mainly provide technical advice to the network of RDS project officers) commented that it was rare to find bats listed in a scheme application, and that this was usually in connection with modifications to farm buildings. The situation may be slightly different in Scotland where species appear to be given greater weight in the supporting evidence to an application, with point scoring for the presence and targeting of approximately 30 habitats prioritised in a local council biodiversity action plan. Pipistrelles, brown long eared and daubenton’s bats were stated to have occurred on various local priority lists and were addressed in scheme applications in these areas.

Table 2 lists example habitat management advice given by agri-environment professionals when asked to explain what recommendations would support bats on the farm. In general, advice focused on the maintenance and creation of foraging habitat.

Hedgerows were universally perceived to be important to bats and aspects of hedgerow management (usually managing for connectivity between other foraging habitat, phasing restoration programs, or creation of tall, bushy hedges) were made twice as often as references to any other habitat. Several interviewees expressed concern over the management of mature trees and deadwood on farms, with RDS co-ordinators commenting that a number of unsuitable applications for tree-work are received each year, and could perhaps be combated by efforts to increase awareness of conservation value and protected species issues. Pollarding mature trees near watercourses was a particularly sensitive issue. Attention to watercourses was addressed mainly in reference to prescriptions to remove stock from these areas, and queries regarding tree planting strategy along watercourses were received. Relatively few references were made directly to the value or management of farm woodland. Pasture tended to be referred to indirectly through issues such as the use of ivermectins, extensive management of grassland generally, and encouraging arable reversion in areas that are predominantly arable. A number of interviewees advocated the strategy of a suite of general measures to improve abundance of insects.

Bat roosts were mentioned in the context of potential building restoration work, when the need to consult a specialist to consider potential impact on protected species was raised. In some regions, grants to facilitate building restoration were now dependent on full consideration to this process, to ensure that protected species were considered more routinely than in previous years. Several advisors perceived building restoration grants to be generally outside their remit, particularly since agri-environment schemes support only buildings restored to agricultural use. Buildings restored to other uses (e.g. as business diversification) were funded by other schemes such as Rural Enterprise Scheme. Pressure to convert agricultural buildings to housing was regarded as a potential negative impact to bats and other wildlife (e.g. barn owls) and was felt to be acute in some areas.

Few advisers listed actions to support bats in hibernation. Integrated management recommendations for bats throughout the year were given only in areas with greater horseshoe bat populations, which were the subject of proactive conservation projects.

Real examples of farm conservation work targeting bats, many supported in part by agri-environment schemes or local funding sources are listed in Table 3.

4 Perceived obstacles to implementing conservation for bats on farmland

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Not all individuals interviewed perceived obstacles to implementing conservation measures targeting bats on farmland. RDS biodiversity co-ordinators / ecologists in particular, believed that scheme agreements could be very flexible, so long as adequate supporting evidence for a measure was provided. Furthermore, several advisors commented that "farmers like bats" and were amenable to incorporating features promoting their conservation.

The most common difficulty encountered (see Box 1) was the poor availability of fine-scale species distribution data, restricting the ability to consider bat requirements at farm or colony level. In the case of rare bats (horseshoe and barbastelle) fine-scale geographic data enabled prioritisation of particular management options within 4 km radius of maternity roosts and hibernacula. Information on other species is patchy, at coarse scale (e.g. 10km 2 not tetrad), or held by other organisations and unavailable if classed as confidential. Some individuals had better access to roost or distribution records, through close associations with local wildlife trusts or bat groups, or in the case of CCW, because the organisation administers both the agri-environment scheme Tir Gofal, and species protection work. When data is available, however, it may be out of date and of questionable relevance. Advisors therefore rely heavily on information obtained from the land manager, since the time taken to gain supporting evidence for an application is limited and does not allow survey for each of the many BAP species in U.K.

Due to their nocturnality, and low profile in scheme information (in comparison to a popular group such as birds), farmers may not always provide such information when habitats and species of interest are considered. Individual advisors had not been able to get funding for species survey, where a rare bat species was suspected, to provide supporting evidence for a management agreement.

Other obstacles mentioned were conflicts of interest with other site objectives or land-manager interests and limitations in what could be achieved by existing agri-environment schemes. A major issue here is that the proportion of farmers participating in agri-environment schemes is relatively small, with uptake being unevenly spread. Farmers outside of agri-environment schemes / local initiatives bear the full-cost of management, so a personal motivation is required to implement conservation-type measures.

Table 2. Recommendations made by agri-environment professionals for habitats they considered to be of particular importance to bats Advisor recommendations for habitat management to support bats No. times cited  Hedgerow and linear feature management (manage for connectivity / obtain 22 tall bushy hedgerows / phase cutting / laying operations over time)  Consider presence of bats in building restoration (requirement to gain 9 specialist advice)  Consider bats in tree work, particularly in pollarding, removal of deadwood 9 or ivy  General measures to encourage invertebrates 8  Fence off and remove livestock from water margins 8  Create grass margins adjacent to linear features 6  Practice more extensive grassland management including management of 6 old pasture  Avoid use of ivermectins 5  Maintain lowland woodland, rides and edges 5  Maintain habitat corridors (i.e. linear features) 4  Revert arable land to grassland and encourage return of livestock 3  Install bat boxes 3  Install ponds 2  Value dung heap as a wildlife resource 1  Consider conversion of military pillboxes to hibernacula 1  Replace standard trees in hedgerows 1  Retain scrub as foraging habitat 1  In vicinity of greater horseshoe roosts follow a series of tailored, integrated measures relating to hedgerow, grassland and stock management according to EN greater horseshoe guidelines Table 3. Examples of conservation measures undertaken by individual farmers that were targeted at bats as part of agri- environment schemes or other local initiatives.

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Examples of farmland conservation work targeting bats Region Example Cambridgeshire / Large farm undertook a FWAG Biodiversity Action Plan and chose to focus Hertfordshire on bats. Riverside management addressed and many bat boxes erected. (This work was not done as part of an agri-environment agreement). Organic farm undertook a FWAG Biodiversity Action Plan and chose to focus on bats. Managed the river corridor, maintained ivy on mature trees. Devon The Greater Horseshoe Bat project. Specific management plan with detailed guidance on grazing regimes tailored to individual farms and proactively targeted at farms within 4km or greater horseshoe roosts. Farms use alternatives to ivermectin stock treatments, avoid rolling and harrowing grass to enhance cockchafer populations and phase hedge management. Winter grazing encouraged near hibernacula. In Exmoor area, barbastelle local radio-tracking information used to support management protocols in ESA management of riparian woodland, hedgerows and wet grassland. Farmers now targeted outside 4km radius of roost where radiotracking provides evidence of habitat use. Gloucestershire Farmer erected bat boxes in a woodland. Boxes checked by advisor in partnership with local bat group In vicinity of greater horseshoe roost, hedgerow restoration policy changed to ensure connectivity of tall, hedgerows at a site where combination of coppicing and hedge laying would otherwise have been adopted since farmer was unaware of bats. Bat boxes erected to raise general awareness of bats. Herefordshire Cider apple farm erected bat boxes, created grass field margins adjacent to boundary features and instigated hedge management sympathetic to bats. Kent Grating dene hole on farmland improved farmer’s Countryside Stewardship application. Countryside Stewardship funded the conversion of a military pillbox to a bat hibernacula as a capital project. Northumberland Farm in stewardship with tall straggly hedges that would previously have been restored purely by coppicing left standard trees in hedgerow restoration and planted corner trees to promote bat foraging. Promoted connectivity of hedgerows between woodland and green lanes. Under stewardship farm created ponds and restored hedges to link these features to an area of grassland restoration. Somerset & Avon Land managers in the Mendips score an extra point in scheme application if measures supporting bats can be incorporated in their proposal. Extra payments are available to farmers who treat stock with benign alternatives to ivermectins or create a coral area for stock treatment. West Midlands Rural Enterprise scheme project to restore River Monnow river quality incorporated consideration for bats when coppicing and pollarding bankside vegetation and trees. Ayrshire & Aran Several intensive dairy farms entering the "White & Wild Scheme" chose to undertake measures targeted at bats including planting and restoration of hedgerows, pond creation and tree planting. Cairngorms Several farms have erected bat boxes. General awareness raising among farmers.

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Box 1. List of factors perceived by agri-environment professionals to impede instigation of conservation measures for bats on farmland. Perceived obstacles to implementing bat conservation measures on farmland Knowledge or awareness  Advisor lacks general knowledge about bats (including identification, and ecology with respect to farmland)  Advisor lacks information on presence of bats or species distribution information at scale appropriate to farm e.g. tetrad level.  Bats have low profile with farmer (e.g. seen infrequently, lack of information targeted at farmer)  Bats are not prioritised in agri-environment schemes e.g. Countryside Stewardship does not list bat species directly etc; regions where no bat species listed on LBAP. Short-comings of agri-environment schemes  Schemes are habitat and landscape focused rather than species focused  Sometimes difficult to incorporate novel projects, particularly if scheme administrators have little experience with such approaches leading to potential scepticism over project value / chances of success.  Cost. Conservation measures funded at different levels. Where measures funded at low levels e.g. 20% of cost, other motivation is necessary to justify cost to land-manager.  Uptake limited to a small proportion of land managers i.e. in some regions <10-15% farmers in Countryside Stewardship.  Reactive rather than proactive approach. Agri-environment schemes are voluntary and except in the case of particular local initiatives there is a lack of opportunity to encourage pre-selected land- managers to apply (i.e. areas of high wildlife value).

 Small farms may find entering agri-environment schemes difficult because they may not wish / be able to propose several different conservation management features. (i.e. farms that wish to focus solely on hedgerow management at a disadvantage although this may be of strategic benefit to wildlife).  Scheme appeal varies across different agricultural sectors i.e. application by farms in marginal areas usually higher than application by lowland intensive farms  Schemes and their scoring systems sometimes felt to be too prescriptive or inflexible  Pressure to process agreements rapidly results in less opportunity to consider individual cases in great detail. Conflicts of interest  Other site interests e.g. scrub removal for benefit of plants / archaeology versus potential foraging habitat  Coppicing / laying hedges may benefit small mammals and stock management by bushing out lower hedge, but must be phased to ensure tall hedgerows also available for bats.  Landowners sometimes perceive bats as "getting in way" of other aims e.g. restoration of old buildings for business diversification.  Pressure to convert derelict buildings into housing / other uses in some regions  With business objectives e.g. use of insecticides, use of ivermectins

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5 Local initiatives and new opportunities

Alternative funding for conservation work on farms is available through some local councils and Forestry Commission (e.g. Farming Connect Scheme in Wales). English Nature and Environment Agency are among organisations that fund specific awareness raising campaigns in agricultural target areas. Some of these projects focus on educational access.

Several advisors outlined the increasing trend of environmental credentials to be considered in farm assurance schemes driven by major buyers (supermarkets). FWAG are currently funded to implement farm biodiversity action plans as part of farm assurance schemes for lettuce and beef. Farmers choose target species from a FWAG list, and implement management plans to enhance local populations. Several businesses have focused on bats under this scheme. "White & Wild" a Wildlife Trusts initiative pays a premium to selected dairy farmers who have committed to following farm conservation plans drawn up by FWAG. A number of "White & Wild" intensive dairy farmers in Ayrshire focused on habitats for pipistrelles.

Local initiatives provide the opportunity for conservation advisers to contact farmers in target areas directly, to raise awareness of particular environmental issues, and or / recruit into agri-environment or other grant schemes. Partnerships for targets such as black grouse, or butterflies have proved successful with the ability to address species / issues individually and maximise local impact by encouraging neighbours or groups of land managers to participate.. The Greater Horseshoe Project in south-west England is a successful example and a similar project is intended for the Somerset / Avon area. Evidence of farmer engagement with these schemes is provided by internet searches, showing a number of businesses using greater horseshoe bats in their product marketing.

A bird initiative, the RSPB’s "Volunteer & Farming Alliance" matches farms with trained volunteers who conduct surveys, to provide information on species presence and use of habitat features. This appears to have been popular with farmers, and were it extended to other species groups might increase the pool of species distribution data available to agri-environment professionals.

The current revision of English agri-environment schemes may provide new opportunities. In the new entry-level scheme, a proposal to restrict hedge cutting to twice in five years may result in wide landscape and attitude changes. Wider promotion of group applications would be beneficial, particularly in relation to flooding schemes, where changes to water table level may impact on neighbours and can only proceed with agreement from all affected parties.

6 Support tools for advisors

Advisors identified a broad range of existing resources that support their work (Box 2). Information supporting bat conservation work was drawn from a broad range of sources, including in house electronic technical knowledge systems, but there was particular emphasis on access to colleagues with specialist knowledge, or to local experts. Some felt the resources available to them were already sufficient. Chief among items identified that might enhance bat conservation (see Table 4) was access to specific fine-scale local information on local species distributions, and more information on bat ecology targeted at agricultural situations. An information leaflet targeted at farmers was expected to raise awareness among land managers.

Box 2. Items identified by agri-environment professionals as resources at their disposal that support them in their work, with a particular focus on resources supporting bat conservation work. Resources utilised by agri-environment professionals Technical electronic DEFRA Magic system, GIS (GENI), RDS intranet, FWAG knowledge base system, BCT website Electronic database containing species information (not bats) Digital camera to personalise field reports Publications Books e.g. JNCC Habitat Management for Bats; EN Batworker’s Guide; district Biodiversity Action Plan booklets, Biodiversity Audit for north-east Scotland. Leaflets e.g. EN Focus on Bats, BCT species leaflets, SNH leaflets, Vincent Wildlife Trust leaflets, FSC bat identification sheet, ADAS, SAC Advisory Note, RSPB Other specialists English Nature, local bat group, county ecologist, RDS technical support team, colleagues in FWAG /

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RDS with specific bat experience or holding bat licence, Environment Agency, local wildlife trust, Bat Conservation Trust Other Evidence collected by agreement holder or partner organisation Local Environmental Records Centre Bat training course (introduction or species conservation licence) FWAG landwise document incorporating four BAPs for bats Adviser’s accumulated personal field experience

Table 4. Items identified by advisors, biodiversity co-ordinators and agri-environment ecologists as resources that would enhance their ability to instigate conservation management for bats on farmland. Resources desired by agri-environment advisors Demand score  Specific fine-scale local information on bat roost locations and species 8 distributions  Information on the ecology of different species including how to identify 7 roosts, how bats use farmland, and potentially harmful farming operations  Introduction to bats training course for agri-environment professionals 5  An information leaflet targeted at farmers 4  Simple guidelines e.g. BAP framework tick sheet for scoring habitat features of 1 value to bats.  Information on local contact points for bat-related issues 1  In Scotland, co-ordinated electronic access to habitat designation information 1 (SSSI, SAC) and protected species occurrence at farm-scale

7 Discussion

This exercise set out to review the evidence for past and continuing impacts of agricultural management on bats, to find out how conservation measures are implemented on farmland, and to identify new opportunities for research and practical implementation. In recognition that much ecological research on bats has been comparatively recent, the review also collated information on several aerial-insect feeding birds which, through some similarities in ecology, might be exposed to similar pressures. Information on eleven British bat species which had been studied in British or European landscapes dominated by agriculture was collated from published research, unpublished data and discussions with researchers and other professionals.

It was notable that the review drew on a smaller pool of data than would have been available for most farmland birds. For most bats we encountered in the order of up to five detailed studies, with further information drawn from surveys examining the activity or distribution of several species or groups. Due to the small pool of studies there is at present little understanding of the range of intraspecific variation in diet, foraging ecology, individual and colony home range and general utilisation of habitat. The new availability of radio-tracking technology for bats is changing previous conceptions. For example, a "woodland species" brown long-eared bat was recorded commuting along drainage ditches in open country, and travelling much greater distances between roosting and foraging sites than had been previously recorded (Moore et al., (submitted)), while another "woodland" species, natterer’s selected arable fields as foraging sites (Aegerter, pers. comm.). There is therefore much scope for developing further research across a range of landscape and management gradients, particularly where this might provide information on behaviour in "favourable" or "impoverished" environments. Relatively little information was available concerning the aerial-insect feeding birds considered appropriate to the review (swallow, house martin, swift, spotted flycatcher). Swallow ecology on farmland was similar in a number of respects to that of some bats (e.g. close association with buildings, aerial trawling strategy, central place foraging, foraging along field boundaries etc.) but conflicts between population trends recorded in Britain and Europe, make the wide-scale impact of agriculture on this species unclear.

Bat studies were located mainly in lowland agricultural landscapes dominated by grass- or mixed farming, with a paucity of information in arable or upland. This is in contrast to the direction taken by farmland bird studies which focused originally on arable / mixed landscapes, and have examined the impacts of intensification in grassland landscapes comparatively recently e.g. (Fuller, 2003). Much farmland bird research occurred in response to population trends revealed in long-term monitoring programmes that separated trends for distinct land-uses (farmland, woodland

64 Project DEFRA title project code etc.). The National Bat Monitoring Programme provides a mechanism for the development of similar indices for bats, with the capacity to improve understanding of the future impacts of agriculture on biodiversity.

In the main, studies focused on determining basic ecology such as the identification of foraging habitat preferences, distances travelled between roost and foraging area and diet rather than on comparing responses to specific aspects of agricultural practice. Exceptions were (Wickramasinghe et al., In press) and Mathews (pers comm.) who compared the activity of bats on organic and conventional farmland, (Moore et al., (submitted)) who compared bat use of recently established farm woodlands with adjacent arable land and (Vaughan et al., 1996) who sampled bat activity upstream and downstream of point pollution sources in order to evaluate the implications of diffuse pollution from agricultural land. There is clearly the opportunity to evaluate the impacts of agri-environment prescriptions, changes in land-use etc. in focused studies but also to benefit from new attitudes toward agri-environment research. (Benton et al., 2003) propose that future research should develop cross-cutting policy frameworks and management solutions to recreate heterogenity in temperate agricultural systems. This argument is a response to the previous research emphasis on the investigation of specific practices in a system where impacts have occurred at multiple temporal and spatial scales.

Relatively few of the papers encountered integrated sampling of bats, habitats and the abundance of insect prey. Those that did tended to find that bats foraged in areas of higher insect density e.g. along stretches of river bordered by trees (Warren et al., 2000), along sheltered stretches of hedgerow (Verboom & Spoelstra, 1999). Studies of grassland birds have found more complex relationships e.g. (Fuller, 2003) in which birds do not always forage where invertebrate densities are highest, presumably because dense vegetation structure impedes access or reduces the efficiency with which prey may be detected or captured. Strictly aerial predators (i.e. a hawking, trawling, or fly-catching strategy) might make simple foraging choices in relation to food abundance, but gleaners or those that take prey from the ground, e.g. (Catto et al., 1996) may experience similar interactions with vegetation structure. Even in the case of predators that capture prey away from vegetation, further research may reveal behaviours that balance prey capture with detrimental consequences such as predation. Predator avoidance has been proposed in explanation of species whose behaviour in open habitats varies at different stages of the night.

In comparison with birds, for which precise visual observations may usually be accrued, current techniques for studying bats (i.e. radiotracking) allow less certainty as to location in relation to habitat. On occasions this is likely to result in interactions between closely located habitats e.g. boundary x crop effects being overlooked. Greater resolution of habitat utilisation, and habitat descriptions that provide more detail on land-use e.g. winter or spring cereal will lead to better understanding of the value of agricultural habitats for bats. Apart from greater horseshoe bats, which have been studied intensively in all life-history stages (breeding, hibernation, foraging etc) the majority of studies examined the distribution of foraging or commuting bats, with few studies providing detailed data on roosting or hibernation requirements on agricultural land. Lack of data on all requirements may result in increased vulnerability at certain stages. In the short term, greater information on roost sites on agricultural land would enable agri-environment advisors to consider broader aspects of management.

In comparison to groups such as ground-beetles and spiders, there was a paucity of ecological information available on the aerial-insects that feature in the diet of bats (Appendix 4). A further difficulty occurs in that the relevance of the data that exist is uncertain due to the coarse identification of prey in bat diet. Insect families often comprise species with widely contrasting life-histories and habitat requirements and generalisations regarding the precise management that would improve prey abundance are therefore more difficult. Of more concern is the lack of information on aerial invertebrate distribution at assemblage level, as studies tended to be of single species or of general abundance, with little information on diversity changes in response to management or habitat. Although bat and aerial-insect feeding birds are unlikely to make distinction between species, (except possibly in relation to prey size) high diversity invertebrate assemblages are likely to provide a greater continuity of food-resource than high abundances of single species. The pragmatic approach of managing for habitat heterogeneity in time and space is likely to apply at least until the requirements of prey groups are better understood. The lack of data on the ecological requirements of moths, and on the distribution of moth assemblages, is a case in point. Example requirements are usually extrapolated from more well-studied butterflies but assumptions of this type are likely to lack precision.

Despite the lack of historical bat monitoring data during the period when greatest agricultural intensification occurred, there is sufficient indirect evidence from data on changes to prey, habitats and recently on bat activity in "intensively" versus "extensively" managed farmland landscapes (Wickramasinghe et al., In press) to suggest that, in common with other farmland wildlife groups, intensification processes were likely to have been detrimental to bat populations. The studies encountered reported a high degree of association between bats and the non-crop elements of farmland

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(woodland, water, hedgerow etc.) and that bats tended to forage in areas with higher insect densities. Operations which reduce the area of non-crop habitats in the landscape, or reduce the quality of such habitats e.g. diffuse pollution of watercourses, reduction of boundary invertebrate fauna through insecticide drift etc are likely to be detrimental to bats. Diversity and abundance of many invertebrate groups e.g. ground beetles and Lepidoptera is usually highest at the field boundary. Measures that target the conservation of species in these areas perform the multiple functions of focusing conservation effort where crop production is poorest, where invertebrate diversity is richest and along linear features where bats are widely reported to commute or forage. Although loss of non-crop habitats such as ponds and hedgerows appears to have halted or reversed, declines in habitat quality remain an issue, e.g. a decline in species richness of hedge bottom plants (including nectar and food-plants for Lepidoptera) during the past decade (Haines-Young et al., 2000).

A further lesson from bird-research that may be pertinent to future approaches to bat research on farmland is the limitation of abundance and activity data in revealing the true impact of management on populations (Gillings & Fuller, 1998). Bird researchers have found that calculation of survival and reproductive success provide better measures of habitat quality since they can be conducted at farm or patch-scale, provide insights into the mechanisms behind population trends, and may be determined over shorter time periods. The current suite of agri-environment schemes provides many prescriptions for habitats that are important to bats (Appendices 5-6). Interestingly few of these were listed to be of particular importance for mammals, and many were developed through bird research. The schemes are habitat focused rather than species focused in the majority of cases, but agri-environment scheme literature could be adapted to raise awareness of potential benefits for mammals. In practice the benefits of schemes are interpreted to land-managers by agri-environment professionals, who were in most cases aware of conservation issues relating to bats. The appearance of bat species on local priority lists tended to raise awareness. The greatest obstacle to instigating conservation measures for bats was lack of species distribution data at a fine-spatial scale. This could potentially be addressed by wider monitoring programmes, and improved data-sharing between appropriate organisations.

66 Fig 1. Proportions of major land-use types on agricultural land in England (DEFRA, 2002)

England Total

Eastern South-east South-west

East Midlands West Midlands

Yorkshire & Humberside North East North-west

Grass/rough grazing Cereals Horticulture

Setaside Other crops/fallow Woodland

All other land

CSG 15 (Rev. 6/02) 67

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